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number 22

June 1995

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EDITORIAL STAFF

OCT * • Richard A. Lancia, Editor

U C STATE UBBAKi Suzanne A. Fischer, Assistant Editor

D&l ftfJH Eloise F. Potter, Production Manager

EDITORIAL BOARD

James W. Hardin

Professor of Botany

North Carolina State University

William M. Palmer

Director of Research and Collections

North Carolina State Museum

of Natural Sciences

Rowland M. Shelley

Curator of Invertebrates

North Carolina State Museum

of Natural Sciences

Robert G. Wolk

Director of Programs

North Carolina State Museum

of Natural Sciences

Brimleyana, the Zoological Journal of the North Carolina State

Museum of Natural Sciences, appears twice yearly in consecutively

numbered issues. Subject matter focuses on systematics, evolution,

zoogeography, ecology, behavior, and paleozoology in the southeastern

United States. Papers stress the results of original empirical field

studies, but synthesizing reviews and papers of significant historical

interest to southeastern zoology are also included. Brief communications

are accepted.

All manuscripts are peer reviewed by specialists in the Southeast

and elsewhere; final acceptability is determined by the Editor. Address

manuscripts and related correspondence to Editor, Brimelyana, North

Carolina State Museum of Natural Sciences, P.O. Box 29555, Raleigh,

NC 27626. Information for contributors appears in the inside back

cover.

Address correspondence pertaining to subscriptions, back issues,

and exchanges to Brimleyana Secretary, North Carolina State Museum

of Natural Sciences, P.O. Box 29555, Raleigh, NC 27626.

In citations please use the full name - Brimleyana.

North Carolina State Museum of Natural Sciences

Betsy Bennett, Director

North Carolina Department of Environment,

Health and Natural Resources

James B. Hunt, Jr., Governor

Jonathan B. Howes, Secretary

CODN BRIMD 7

ISSN 0193-4406

Aspects of the Feeding Ecology of the

Little Grass Frog, Pseudacris ocularis

(Anura: Hylidae)

Jeremy L. Marshall1 and Carlos D. Camp

Department of Biology, Piedmont College

Demorest, Georgia 30535

ABSTRACT— We report on the foods of the little grass frog,

Pseudacris ocularis, from Georgia. Fifty specimens were collected

from two isolated wetlands located in Evans and Grady counties,

Georgia, during late spring and summer 1993. Analysis of stomach

contents determined that the most abundant food items were

small arthropods associated with leaf litter and soil. Almost

50% of the food items were collembolans, followed by hymenopterans

(17%), acarines (9%), homopterans (8%), and coleopterans (8%).

We compared foods of adult males with those of newly meta-

morphosed juveniles collected at the same time from the Grady

County site. Juvenile frogs ate more individual food items and

a greater diversity of prey species than did adult males. This

difference could be due to adult Pseudacris selecting larger,

more profitable prey than juveniles select. Lower feeding activity

exhibited by breeding males might also be a contributing factor.

Little is known about the feeding ecology of many amphibians,

especially intraspecific variability in foods and foraging (Duellman

and Trueb 1986). Variation in dietary preferences among population

subgroups (e.g., breeding males, non-breeding females, subadults,

juveniles, larvae, etc.) has been reported to reflect differences in

habitat preference (Lamb 1984), gape (Toft 1980), developmental

condition (Brophy 1980, Davie 1991), and other factors.

The little grass frog, Pseudacris ocularis (Bosc and Daudin), is

the smallest North American anuran (Conant and Collins 1991). It

occurs in a wide variety of ephemeral and semi-permanent wetlands

in the southeastern Coastal Plain and favors grassy areas in and around

cypress ponds and similar sites (Harper 1939, Mount 1975). In spite

of its relative abundance in many of these areas, virtually nothing

is known of the feeding ecology of this frog. The purpose of our

study was to describe the diet of P. ocularis and to investigate any

potential differences between the feeding of adult frogs and newly

metamorphosed juveniles.

1 Present address: Biology Department, University of Mississippi, University,

Mississippi 38677.

Brimleyana 22:1-7, June 1995 1

Jeremy L. Marshall and Carlos D. Camp

MATERIALS AND METHODS

We collected 50 P. ocularis for stomach analysis from two

localities during May-July 1993. Both sites were ephemeral wetlands

in the lower Coastal Plain of Georgia. The first site, located in Grady

County, was dominated by black gum (Nyssa sylvatica) and was situ-

ated in a low pine flatwood having a canopy of slash pine (Pinus

elliottii) and an understory of saw palmetto (Serenoa repens) and

gallberry (Ilex glabra). The second, in Evans County, was a dome of

pond cypress (Taxodium ascendens) surrounded by sandhills domi-

nated by longleaf pine (P. palustris) and turkey oak (Quercus laevis).

Areas similar to both sites were described in detail by Wharton (1978).

After collection, all specimens were preserved in 10% formalin,

and stored in 35% isopropanol. Each frog was measured for snout-

vent length (SVL) and dissected for stomach analysis. Individual food

items were counted and identified. Because prey items were too small

to use volumetric displacement, relative importance of prey was de-

termined by comparing each individual prey item to a paper grid and

visually estimating the number of grid squares occupied (Camp and

Bozeman 1981).

Twenty of the Grady County frogs were collected between 2200

and 2400 EDT on 5 June. This sample consisted of 10 mature males

and 10 juveniles that had just completed metamorphosis. We used

this sample to make comparisons between feeding of adults and juve-

niles. Because we did not independently test for prey availability,

other collections were not used for comparisons because of possible

complications arising from temporal or between-site differences in

available prey items. In addition, although adult females were in-

cluded in these samples, small numbers (n = 4) precluded between-

sex comparisons. Correlation between the number of prey items eaten

and body size was tested using the procedure described by Zar

(1984). A comparison of diversity between adult male and juvenile

prey species was made using the Shannon-Wiener Index of Diversity

(H') (Zar 1984).

RESULTS

One hundred-forty individual prey items were identified and

consisted entirely of arthropods, mainly insects (Table 1). Springtails

(Collembola) were the most numerous group, making up 47% of the

food items eaten and found in 56% of the stomachs. Because they are

so small, however, they contributed less than 20% of the area occu-

pied by all prey items. Hymenopterans, especially ants (Formicidae)

Little Grass Frog Feeding Ecology

Table 1. Stomach contents of 50 Pseudacris ocularis from the Coastal

Plain of Georgia. Unless otherwise indicated, the smallest taxon in

each order is represented by a single species; "i" repreesnts immature

instars. Numbers (n) for higher taxa also include unidentified food

items.

Percentage of

Food Item

INSECTA

Collembola

Isotomidae

Poduridae

Sminthuridae

Coleoptera

Carabidae

Cleridae

Coccinellidae

Staphylinidae (3)

larvae

Dictyoptera

Blattidae

Diptera (larvae) (3)

Homoptera

Delphacidae (i)

Hymenoptera

Diapriidae

Dryinidae

Encyrtidae

Evaniidae

Formicidae

Scelionidae

Orthoptera

Acrididae (i)

Phasmida

Phasmatidae

Siphonaptera

Thysanoptera

Phlaeothripidae

ARACHNIDA

ACARINA

Mesostismata

Oribatei (2)

Araneida

Anyphaenidae

Palpigradi

(n) Total Number Total Area Frequency

122

66

26

2

37

11

1

5

1

2

3

11

10

24

1

2

7

2

1

18

13

1

12

4

1

87.1

47.1

18.6

1.4

26.4

7.9

0.7

3.6

0.7

1.4

2.1

7.9

7.1

17.1

0.7

1.4

5.0

1.4

0.7

12.9

9.3

0.7

8.6

2.9

0.7

90.1

19.5

5.6

0.5

13.4

9.2

1.6

0.3

1.1

4.3

0.4

19.1

1.2

11.1

10.9

14.6

2.2

0.8

0.1

6.4

1.2

1.9

4.8

9.6

0.1

9.9

5.5

0.1

5.4

4.2

1.6

0.2

82.0

56.0

20.0

4.0

26.0

18.0

2.0

6.0

2.0

6.0

16.0

14.0

32.0

2.0

6.0

4.0

2.0

22.0

16.0

2.0

14.0

6.0

2.0

Jeremy L. Marshall and Carlos D. Camp

and parasitic wasps (Scelionidae), were the second most important

group, making up 17% numerically and occurring in 32% of the

stomachs. Hymenopterans made up 15% of the relative area. Other

important insect groups represented were coleopterans, particularly

rove beetles (Staphylinidae), making up 8% of numbers, 9% of area,

and occurring in 19% of the stomachs, and delphacid homopterans

(7, 11, and 14%, respectively). Although found only occasionally,

relatively large roaches (Dictyoptera) and walking sticks (Phasmida)

made up a considerable amount of the total quantity of food eaten

(19 and 10% of total area, respectively). The only non-insect food

items found were arachnids. These consisted primarily of mites

(Acarina), which made up 9% numerically, 6% of the area, and occurr-

ed in 16% of stomachs.

Juvenile frogs from the 5 June, Grady County sample had a

mean SVL of 8.80 mm with a standard error (SE) of 0.21 mm. Adult

males from this sample had a mean SVL of 14.87 mm and a SE of

0.23 mm. Food items eaten by these frogs are shown in Table 2. The

Shannon-Wiener Index for juvenile prey species diversity (//' = 1.062)

was significantly larger than that for adults {H' = 0.739; t = 3.27, df

= 45, P < 0.01). There was a negative correlation between number

of food items eaten and frog size (r2 = 0.25, t = 2.18, df = 14, P <

0.05).

DISCUSSION

Pseudacris ocularis is commonly found on lower tree trunks and

foliage up to a height of 1 m or more (Harper 1939); males prefer

these sites as calling perches (Harper 1939, Mount 1975). However,

the majority of food items we found were arthropods that are associ-

ated with leaf litter and/or soil (e.g., springtails, mites, dipteran lar-

vae, staphylinids, ants, thrips, palpigrades, etc.). In addition, we found

a large number of frogs on the ground, particularly during daytime

collections. It is apparent, then, that P. ocularis spends a consider-

able amount of its foraging time on the ground.

According to optimal foraging theory (Pyke et al. 1977, Krebs

1978), a predator should choose prey that represent the greatest net

energy gain and forage in areas where profitable prey are most fre-

quently encountered. Considering the small size of these frogs, small

abundant leaf litter arthropods such as springtails and mites might

represent a relatively stable, predictable source of profitable prey.

However, amphibians might find larger arthropod prey to be more

profitable than small ones due to a proportionately smaller exoskel-

eton (Jaeger and Barnard 1981). Therefore, little grass frogs should

Little Grass Frog Feeding Ecology

Table 2. Stomach contents of 10 juvenile and 10 adult P. ocularis collected

5 June 1993, Grady County, Georgia; "i" represents immature instars;

* indicates two species represented. Numbers for higher taxa also include

unidentified food items.

feed more on larger prey when available. Our data would, in part,

appear to confirm this hypothesis. For instance, relatively large im-

mature delphacids made up < 7% of total food items. However, in the

5 June, Grady County sample of adults (Table 2), delphacids made up

23%, indicating these food items were probably more available at

that time, although we do not have independent confirmation of prey

abundance.

Newly metamorphosed P. ocularis ate more individual food

items and a greater diversity of prey species than did adult males.

Two factors may explain these results. First, there may be an ontogenetic

shift in foraging strategy during post-metamorphic growth of P. ocularis.

Such a shift has been inferred in P. triseriata (Christian 1982) where

Jeremy L. Marshall and Carlos D. Camp

adults select more optimal (i.e., large) prey than do juveniles, which

indiscriminately feed on prey they encounter. This may be the result

of larger animals being able to choose from a greater range of prey

sizes, whereas smaller individuals are largely restricted to small prey,

as is apparent in P. crucifer (Oplinger 1967). This shift would account

for the lower diversity of prey species taken by adult P. ocularis.

Second, the adult sample used in our comparisons consisted entirely

of males. Several authors have reported a sharp decline in feeding

activity by adult male frogs during the breeding season (Jenssen and

Klimstra 1966, Lamb 1984). The males in our study were not breeding

(the pond was completely dry) and only sporadically calling at the

time of collection (5 June), although breeding had been previously

observed at this site in March. Mount (1975), however, reported breeding

congregations of P. ocularis as late as 29 July in nearby Houston

County, Alabama, and Harper (1939) recorded vigorous chorusing in

the Okefenokee during August and September. Therefore, since P.

ocularis does breed throughout the summer, we cannot rule out the

possibility of lower feeding activity in adult males during the time of

our collections.

ACKNOWLEDGMENTS— Thanks are extended to Tyler Lee

(Cairo, Georgia) and Ben Cash (Georgia Southern University) for

help in collecting specimens, as well as Rob Wainberg (Piedmont

College) for assistance in identifying prey items. In addition, Rob

Wainberg and Rick Austin (University of Mississippi) critically read

the manuscript.

LITERATURE CITED

Brophy, T. E. 1980. Food habits of sympatric larval Ambystoma

tigrinum and Notophthalmus viridescens. Journal of Herpetology

14:1-6.

Camp, C. D., and L. L. Bozeman. 1981. Foods of two species of

Plethodon (Caudata: Plethodontidae) from Georgia and Alabama.

Brimleyana 6:163-166.

Christian, K. A. 1982. Changes in the food niche during post-

metamorphic ontogeny of the frog Pseudacris triseriata. Copeia

1982:73-80.

Conant, R., and J. T. Collins. 1991. Reptiles and amphibians of

eastern/central North America. Houghton-Mifflin, Boston, Massa-

chusetts.

Little Grass Frog Feeding Ecology

Davie, R. D. 1991. Ontogenetic shift in diet of Desmognathus quadramacula-

tus. Journal of Herpetology 25:108-111.

Duellman, W. E., and L. Trueb. 1986. Biology of amphibians. McGraw/

Hill, New York, New York.

Harper, F. 1939. Distribution, taxonomy, nomenclature and habits of

the little grass frog {Hyla ocularis). American Midland Naturalist

22:134-149.

Jaeger, R. G., and D. E. Barnard. 1981. Foraging tactics of a

terrestrial salamander: choice of diet in structurally simple envi-

ronments. American Naturalist 117:639-664.

Jenssen, T. A., and W. D. Klimstra. 1966. Food habits of the green

frog, Rana clamitans, in southern Illinois. American Midland Naturalist

76:169-182.

Krebs, J. R. 1978. Optimal foraging: decision rules for predators

Pages 23-63 in Behavioral ecology: an evolutionary approach

(J. R. Krebs and N. B. Davies, editors). Sinauer Publications,

Sunderland, Massachusetts.

Lamb, T. 1984. The influence of sex and breeding condition on

microhabitat selection and diet in the pig frog Rana grylio. American

Midland Naturalist 111:311-318.

Mount, R. H. 1975. The reptiles and amphibians of Alabama. Ala-

bama Agricultural Experiment Station, Auburn University, Auburn,

Alabama.

Oplinger, C. S. 1967. Food habits and feeding activity of recently

transformed and adult Hyla crucifer crucifer Wied. Herpetologica

23:209-217.

Pyke, G. H., H. R. Pulliam, and E. L. Charnov. 1977. Optimal

foraging: a selective review of theory and tests. Quarterly Re-

view of Biology 52:137-154.

Toft, C. A. 1980. Feeding ecology of thirteen syntopic species of

anurans in a seasonal tropical environment. Oecologia 45:131-141.

Wharton, C. H. 1978. The natural environments of Georgia. Georgia

Department of Natural Resources, Atlanta.

Zar, J. H. 1984. Biostatistical analysis, Second edition. Prentice-

Hall, Englewood Cliffs, New Jersey.

Received 28 June 1994

Accepted 10 November 1994

Digitized by the Internet Archive

in 2013

http://archive.org/details/brimleyana19nort_11

Corrections of Records of Occurrence

of Peromyscus polionotus (Wagner) and P. gossypinus

(LeConte) (Rodentia: Muridae) in the Blue Ridge

Province of Georgia

Joshua Laerm and James L. Boone

Museum of Natural History and Institute of Ecology

University of Georgia, Athens 30602

ABSTRACT — Reexamination of marginal records of Peromyscus

polionotus and P. gossypinus previously reported from the Blue

Ridge Province of Georgia indicate specimens were misidentified.

Neither species occurs in the Blue Ridge Province. The distribution

of P. polionotus is restricted to south of a line from Greenville

and Spartanburg counties, South Carolina, southeast to Clarke,

County, Georgia and west to Dawson and Cherokee counties,

Georgia. The distributional limit of P. gossypinus is south of

a line from Lincoln and Wilkes counties, Georgia west to Dekalb

and Fulton counties and then west and north to Polk, Floyd,

and Dade counties.

Mice of the genus Peromyscus Gloger are among the most

common and broadly studied small mammals in North America. Yet

they are often difficult to distinguish on the basis of traditional

morphological features, and consequently limits to their distribution

are difficult to delineate. This is particularly true within Osgood's

(1909) P. leucopus and P. maniculatus species groups (Hooper 1968,

Laerm and Boone 1994). Frequently biologists depend upon range

maps to rule out certain species; nevertheless, the correct identifica-

tion of a taxon should be based upon morphological characteristics.

Numerous regional studies have been undertaken to provide

mensural discrimination between species of Peromyscus (Choate 1973,

Linzey et al. 1976, Choate et al. 1979, Stromberg 1979, Engstrom et

al. 1982, Feldhamer et al. 1983, McDaniel et al. 1983). Laerm and

Boone (1994) recently used discriminant analysis to maximally

distinguish between the four Peromyscus species that occur in the

southeastern United States: P. gossypinus, leucopus, maniculatus, and

polionotus. Based upon reexamination and correct identification of

specimens representing marginal records of P. gossypinus and P.

polionotus with this discriminant analysis model, we questioned the

accuracy of existing range maps and marginal records for P.

polionotus (Wagner) and P. gossypinus (LeConte).

Brimleyana 22:9-14, June 1995

10 Joshua Laerm and James L. Boone

Peromyscus polionotus — A black-eyed, white Peromyscus was

reported from Tallulah Falls, Rabun County, Georgia by Dice (1934:246)

who identified the specimen as P. polionotus polionotus. This specimen

was noteworthy because the pelage was entirely white, and the feet,

toes, and nails lacked any pigment. However, as Dice noted, the eyes,

eye-ring, and outer parts of the ears were dark. Tallulah Falls, in the

Tallulah River Gorge, is at the southern edge of the Blue Ridge

Province in Georgia. While not specifically indicated by Dice, this

would have been the northernmost record of the species, from a

locality well outside the previously described range (Osgood 1909)

and in habitat from which the species had never been reported. Typically,

P. polionotus is restricted to sandy soils and does not occur north of

the middle Piedmont of Georgia.

Schwartz (1954) revised the Peromyscus polionotus complex and

described several new subspecies. He provided external and cranial

measurements of the six subspecies of polionotus he recognized and

referred populations of polionotus in the northern portion of Georgia

and South Carolina (essentially north of the Fall Line) to P. p. colemani.

Schwartz (1954:568) commented on the Dice specimen which he ". . .

presumed, on geographical grounds, to be assignable to P. p. colemani"

Had Schwartz actually examined the Dice specimen, he probably would

not have referred it to P. polionotus. However, Schwartz did not

include the specimen in his mensural analysis. Thus, following Dice

(1934) and Schwartz (1954), Hall and Kelson (1959), Golley (1962),

and Hall (1981) continued to include the specimen as a marginal

record for the species.

We questioned the identification of the Dice specimen because

(1) it was collected in quartzite sheer rock walls and talus of the

Tallulah River Gorge in the Blue Ridge Province and (2) at a locality

some 100 km north of any other specimen record. We compared our

measurements of the Dice specimen (University of Michigan Museum

of Zoology 68496) to measurements provided by Schwartz (1954) for

P. p. colemani and subjected it to our discriminant function (Laerm

and Boone 1994). For 8 of the 11 characters examined by Schwartz

(1954:565), the Dice specimen was larger than the range of the com-

parable measurements made on 11 P. p. colemani by Schwartz.

Visual comparison of means and ranges of measurements of 110

P. polionotus and 107 P. leucopus, which were used to develop a

discriminant function for mensural discrimination between these and

other southeastern Peromyscus spp. (measurements provided in Laerm

and Boone 1994), to the Dice specimen indicates that it generally

falls inside the range of measurements available for both polionotus

Occurrence of Peromyscus

11

and leucopus. Our discriminant analysis, however, strongly suggest the

Dice specimen to be P. leucopus (P = 0.939).

Concern over the correct identification of other northern

marginal range records of P. polionotus in Georgia, South Carolina,

and Alabama (see Schwartz 1954, Golley 1966, Wolfe and Rogers

1969, Hall 1981) prompted an examination of these specimens as

well. We are satisfied that specimens reported from Jackson County,

in extreme northeastern Alabama (Schwartz 1954, Hall and Kelson

1959, Hall 1981) are P. polionotus. Similarly, specimens from Greenville

and Spartanburg counties South Carolina referred to by Schwartz

(1954), Golley (1966), and Hall (1981) are correctly identified as P.

polionotus.

Review of currently available distributional records for P.

polionotus in Georgia now indicates the northernmost limit of its

range should be amended to extend from Spartanburg and Greenville

counties, South Carolina, southwest to Clarke County, Georgia, and

west to Dawson and Cherokee counties, Georgia. Unfortunately, there

are no records available for the Ridge and Valley or Cumberland

Plateau provinces of Georgia west of Dawson and Cherokee counties

(Fig. 1). The next records to the northwest are in Jackson County,

Alabama (Hall 1981).

Fig. 1 Distribution of Peromyscus gossypinus (a) and P. polionotus

(b) in Georgia. Dots represent northernmost distributional records only.

12 Joshua Laerm and James L. Boone

Peromyscus gossypinus — Hall and Kelson (1959), apparently

following Osgood (1909), indicated the range of P. gossypinus in

Georgia to be restricted to the Coastal Plain. At that time no speci-

mens were known from the Piedmont, Blue Ridge, Ridge and Valley,

or Cumberland Plateau provinces. Golley (1962) published new dis-

tributional records for Georgia and indicated (Golley 1962:124)

specimens occurring in the southeastern portion of the Piedmont

(including Columbia, Lincoln, McDuffie, and Wilkes counties),

Ridge and Valley (Floyd, Gordon, and Polk counties), Cumberland

Plateau (Dade County), extreme upper Piedmont (Habesham County)

as well as the Blue Ridge Province (Rabun County). Subsequently,

Wolfe and Linzey (1977) mapped the distribution of the species,

apparently following Hall and Kelson (1959). Wolfe and Linzey (1977)

do not cite Golley (1962); however, Hall (1981) does cite the Golley

(1962) records and maps them accordingly, indicating their range

extends into the extreme northeastern Piedmont and Blue Ridge

provinces.

We used our discriminant function to examine virtually all P.

gossypinus records from Georgia. Specimens from Dade County on

the Cumberland Plateau and those from Floyd and Polk counties in

the Ridge and Valley are P. gossypinus as noted by Golley (1962).

We were unable to locate any museum specimens referred to by

Golley (1962) from Habersham or Rabun counties either in University

of Georgia Museum of Natural History collections or those of all

other North American mammal collections housing specimens from

Georgia. Examination of skin tags and records has not indicated any

P. gossypinus from those counties to have been re-identified and/or

relabeled. We are confident that the specimens of P. gossypinus from

Ruban and Habersham counties in Golley (1962) were erroneously

mapped because, in his verbal description of the range, Golley (1962:128)

indicated P. gossypinus to be found "... on the coastal plain, but

extending into the Piedmont on the eastern margin of the state and

into the ridge and valley province on the west." He makes no mention

of any records in the extreme northern Piedmont or Blue Ridge. Thus,

the range of P. gossypinus in Georgia should be amended to extend

across the middle portion of the Piedmont from Lincoln and Wilkes

counties west to Dekalb and Fulton counties and then west and north

into Polk, Floyd, and Dade counties in the Ridge and Valley and

Cumberland Plateau (Fig. 1). This is essentially the range as previously

depicted by Hall and Kelson (1959) and Wolfe and Linzey (1977).

Occurrence of Peromyscus 13

LITERATURE CITED

Choate, J. R. 1973. Identification and recent distribution of white-

footed mice {Peromyscus) in New England. Journal of Mammalogy

54:41-49.

Choate, J. R., R. C. Dowler, and J. E. Krause. 1979. Mensural

discrimination between Peromyscus leucopus and P. maniculatus

(Rodentia) in Kansas. Southwestern Naturalist 24:249-258.

Dice, L. R. 1934. A black-eyed white Peromyscus. Journal of Mammalogy

15:246.

Engstrom, M. D., D. J. Schmidly, and P. K. Fox. 1982. Nongeographic

variation and discrimination of species within the Peromyscus leucopus

species group (Mammalia: Cricetinae) in eastern Texas. Texas Journal

of Science 34:149-162.

Feldhamer, G. A., J. E. Gates, and J. H. Howard. 1983. Field iden-

tification of Peromyscus maniculatus and P. leucopus in Maryland:

Reliability of morphological characters. Acta Theriologica 28:417-423.

Golley, F. B. 1962. Mammals of Georgia: A study of their distribution

and functional role in the ecosystem. University of Georgia Press,

Athens.

Golley, F. B. 1966. South Carolina mammals. The Charleston Museum,

Charleston, South Carolina.

Hall, E. R. 1981. The Mammals of North America. Second edition.

John Wiley and Sons, New York, New York.

Hall, E. R., and K. R. Kelson. 1959. The Mammals of North America.

Ronald Press, New York, New York.

Hooper, E. T. 1968. Classification. Pages 27-74 in Biology of

Peromyscus (Rodentia) (J. A. King, editor). Special Publication

of the American Society of Mammalogists 2:1-593.

Laerm, J., and J. L. Boone. 1994. Mensural discrimination of four

species of Peromyscus (Rodentia: Muridae) in the southeastern

United States. Brimleyana 21:107-124.

Linzey, A. V., D. W. Linzey, and S. E. Perkins, Jr. 1976. The

Peromyscus leucopus group in Alabama. Journal of the Alabama

Academy of Science 47:109-113.

McDaniel, V. R., R. Tumlison, and P. McLarty. 1983. Mensural

discrimination of the skulls of Arkansas Peromyscus. Proceedings

of the Arkansas Academy of Science 37:50-53.

Osgood, W. H. 1909. Revision of the mice of the American genus

Peromyscus. North American Fauna 28:1-285.

Schwartz, A. 1954. Oldfield mice, Peromyscus polionotus, of South

Carolina. Journal of Mammalogy 35:561-569.

Stromberg, M. R. 1979. Field identification of Peromyscus leucopus

and P. maniculatus with discriminant analysis. Wisconsin Academy

of Science, Art and Letters 67:159-164.

14 Joshua Laerm and James L. Boone

Wolfe, J. L., and A. V. Linzey. 1977. Peromyscus gossypinus. Mammalian

Species 70:1-5.

Wolfe, J. L., and D. T. Rogers. 1969. Oldfield mammals in western

Alabama. Journal of Mammalogy 50:609-612.

Received 25 July 1994

Accepted 28 November 1994

The Masked Shrew, Sorex cinereus

(Insectivora: Soricidae), and Red-backed Vole,

Clethrionomys gapperi (Rodentia: Muridae), in the

Blue Ridge Province of South Carolina

Joshua Laerm

Museum of Natural History and Institute of Ecology

University of Georgia, Athens, Georgia 30602

Eric Brown, Michael A. Menzel

Daniel B. Warnell School of Forest Resources

University of Georgia, Athens, Georgia 30602

Amanda Wotjalik

Museum of Natural History

University of Georgia, Athens, Georgia 30602

William Mark Ford

Daniel B. Warnell School of Forest Resources

University of Georgia, Athens, Georgia 30602

AND

Mary Strayer

Nongame and Heritage Trust Program,

South Carolina Wildlife and Marine Resources Department

P.O. Box 1806, Clemson, South Carolina 29633

ABSTRACT — The first records of Sorex cinereus Kerr are documented

from South Carolina. Pitfall surveys throughout the Blue Ridge

Province resulted in captures from two localities in markedly

mesic, relict, boreal habitats. Additional records of Clethironomys

gapperi (Vigors) were documented including the most southeastern

record. Both S. cinereus and C. gapperi are rare in South

Carolina, largely because of limited areas of appropriate habitats.

The masked shrew, Sorex cinereus Kerr, has the largest range

and exhibits the greatest geographic variation of any North American

Sorex (Hall 1981, Junge and Hoffmann 1981, van Zyll de Jong and

Kirkland 1989). It ranges throughout the transcontinental coniferous

forests from the Canadian Arctic south to the extreme northern por-

tions of the United States with significant extensions south into the

Brimleyana 22:15-21, June 1995 15

16 Joshua Laerm, et al.

montane forests of the Rocky and Appalachian mountains. In the

southeastern United States including Virginia and West Virginia, eastern

Kentucky and Tennessee, North Carolina and Georgia, S. cinereus is

restricted primarily to high elevation montane communities of the

Appalachian Highlands (van Zyll de Jong and Kirkland 1989). To

date, however, there have been no records from South Carolina

(Golley 1966, Mengak et al. 1987).

Previously, the southernmost records of S. cinereus have been

reported from Georgia based upon three specimens reported by

Wharton (1968) from Beech Creek near its confluence with the Talulla

River, Towns County, at an elevation of 807 m. More recently, Ford

et al. (In press) have reported S. cinereus from numerous, widely

scattered localities throughout the Blue Ridge Province of Georgia,

including localities in close proximity to the South Carolina state

line. Similarly, S. cinereus has been reported from several Blue Ridge

Province counties in North Carolina (Polk, Henderson, Transylvania,

Jackson, Macon, and Clay) which are contiguous to South Carolina

(Lee et al. 1982 and unpublished University of Georgia, Museum of

Natural History records). Because S. cinereus is known to occur in

immediately adjacent areas of Georgia and North Carolina and

because seemingly appropriate areas of high elevation habitat exist in

the Blue Ridge Province of South Carolina, we surveyed the moun-

tainous portions of Oconee, Pickens, and Greenville counties specifi-

cally for S. cinereus.

METHODS

From 23 January to 1 May 1994 pitfall trap surveys were

conducted throughout the Blue Ridge Province of extreme northwest-

ern South Carolina including, from east to west, portions of

Greenville, Pickens, and Oconee counties. We totalled 14,000 trap

nights at 17 individual sites. At each site twenty, 32-ounce, plastic

containers (14-cm lip diameter and 17-cm depth) were placed below

ground level adjacent to forest floor debris including stumps, fallen

logs, rocks, etc, for a minimum of 60 days. Approximately 0.14L of

preservative was placed in the bottom of each pitfall. General habitat

features, including dominant overstory and understory vegetation,

aspect, and approximate stand age, of each site were recorded and

elevations estimated from topographic maps. Traps were checked on

a biweekly basis. Specimens were preserved in alcohol for subsequent

reproductive and gut content analysis. Standard body measurements

were taken, and skulls were prepared for confirmation of identifica-

Masked Shrew and Red-backed Vole 17

tion. All specimens were reposited in the mammal collections of the

University of Georgia Museum of Natural History.

RESULTS AND DISCUSSION

We recovered 15 S. cinereus at two of 17 Blue Ridge Province

sites. Both S. cinereus localities were in the northwestern portion of

Oconee County. Seven individuals were recovered from the grounds

of the Walhala Fish Hatchery in a hemlock (Tsuga canadensis) and

rhododendron (Rhododendron maximum) streamside community which

grades upslope into a yellow poplar (Liriodendron tulipifera), mixed

oak (Quercus spp.), hickory (Carya spp.), and white pine (Pinus strobus)

community. Elevation was approximately 760 m. The second S.

cinereus locality (eight captures) was approximately 1.3 km east of

the Walhala Fish Hatchery site in a moderate to mesic mixed oak

and yellow poplar hardwood community at approximately 800 m.

Sorex cinereus was the dominant small mammal recovered in

the Walhala Fish Hatchery site. Fifteen small mammals were recov-

ered in 1,960 trap nights: seven S. cinereus, two S. fumeus, one Sorex

hoyi, one Blarina brevicauda, two Peromyscus maniculatus, and one

Clethrionomys gapperi. The recovery of S. cinereus was fairly evenly

distributed over the trapping period with one or two captured during

each sampling period.

At the second site, also with 1,960 trap nights, 12 S. fumeus,

four S. hoyi, two Peromyscus leucopus, one P. maniculatus, two Blarina

brevicauda, and one Clethrionomys gapperi were captured in addition

to the eight S. cinereus. Here all the cinereus were captured between

20 March and 3 April; six of which were taken in a single pitfall trap

beneath a large, heavily rotted log.

The breadth and intensity of our collection efforts indicate a

restricted distribution of S. cinereus in South Carolina. Sorex cinereus

is regarded as having a boreomontane distribution (Junge and

Hoffman 1981). In the southern Appalachians it has been documented

by Odum (1949), Johnston (1967), Gentry et al. (1968), Linzey and

Linzey (1971), Whitaker et al. (1975), and Lee et al. (1982) in west-

ern North Carolina; Conaway and Howell (1953), Smith et al. (1974),

and Harvey et al. (1991, 1992), in the mountainous regions of eastern

Tennessee; and Pagels and Tate (1976), Pagels and Handley (1989),

Pagels (1991), Kalko and Handley (1993), and Pagels et al. (1994) in

southwestern Virginia. It has not been recorded from elevations be-

low 610 m in southwestern Virginia (Pagels and Handley 1989) or

North Carolina (Linzey and Linzey 1971, Lee et al. 1982). Similarly,

18 Joshua Laerm, et al.

in Georgia, S. cinereus is restricted to high elevation (790-1,370 m)

in markedly mesic habitats with northern affinities (Ford et al. 1994).

Kirkland (1985, 1991) indicated that soricids, in general, are

most diverse in regions characterized by cool moist forests, possibly

by supporting an abundant, stable, and diverse soil invertebrate fauna

upon which shrews depend. Pagels et al. (1994) have shown that the

presence of S. cinereus was significantly correlated with soil moisture

holding capacity and total understory vegetation, and that habitat features

that promote shaded, moist habitats were particularly important in

relict, boreal forest habitats throughout the southern Appalachians.

Although considerable areas of the Blue Ridge Province in South

Carolina meet or exceed the minimum elevations at which S. cinereus

is found elsewhere in the southern Appalachians, boreomontane

habitats are limited there. At the southern limit of the Appalachian

Mountains, much of the mountain habitat in South Carolina is charac-

terized by south-facing aspects with more xeric, mixed oak and pine

communities. Similar xeric south-facing or ridgeline habitats in Geor-

gia yielded few, if any, S. cinereus in recent studies (Ford et al.

1994). In Georgia we encountered S. cinereus primarily at very high

elevations (over 1200 m) or in rich, moist, streamside communities

dominated by hemlock and rhododendron on the Rabun Bald Massif.

West of the Little Tennessee River in Georgia, S. cinereus is

restricted to higher (over 1000 m) elevations, and then they only

occur in restricted habitats with marked northern affinities such as

those described by Wharton (1968).

Our collection site at the Walhala Fish Hatchery is located in

a relatively narrow, steep-walled gorge of the East Fork of the Chat-

tooga River. Wharton (1977) noted that similar streamside forest

communities on the Georgia side of the Chattooga were kept cool and

moist due to complete shading by the hemlock overstory and

rhododendron shrub layer as well as by steep-walled gorges. He noted

that such areas were refugia of more typical northern forest commu-

nities. The Walhala Fish Hatchery, and its associated upslope north-

ern aspect cove hardwood habitat, might represent a limited finger

or refugia in South Carolina for boreal species such as S. cinereus.

The region of the Walhala Fish Hatchery is one of the few sites in

South Carolina that has yielded other small mammals with a typical

boreal distribution including Clethrionomys gapperi (Pivorun et al.

1984) and Peromyscus maniculatus. Other high elevation sites in the

Blue Ridge Province including Sassafrass Mountain, Jones Gap at

Caesar's Head State Park, Saluda Mountain and Hogback Mountain

were trapped but yielded no S. cinereus. Peromyscus maniculatus has

Masked Shrew and Red-backed Vole 19

been recorded at many of these sites (Golley 1966), and we recovered

several specimens at most of these localities. However, Clethriono-

mys gapperi was not reported beyond the Walhalla Fish Hatchery

site until we recovered one in the region of Sassafrass Mountain

(Pickens County, US Hwy 178, 7.4 m north of State Hwy 11). This is

most southeastern record for the species and, like S. cinereus, it

apparently has a very limited distribution in South Carolina.

ACKNOWLEDGMENTS— We appreciate the field assistance of

numerous persons including E. Mitchell, J. Kimbrell, N. Lautenschlager,

B. Smith, L. Grassman, and S. Chalmers. Collecting activities were

conducted under South Carolina Permit #0016-94. We also thank

biologists with the Walhala Fish Hatchery, Sumpter National Forest,

and Caesar's Head State Park for permission to conduct surveys on

United States Forest Service and South Carolina State Park lands.

LITERATURE CITED

Conaway, C. H., and J. C. Howell. 1953. Observations on the mammals

of Johnston and Carter counties, Tennessee and Avery County,

Virginia. Journal of the Tennessee Academy of Science 28:53-61.

Ford, W. M., J. Laerm, D. C. Weinand, and K. G. Barker. 1994.

Small mammal surveys in the Chatahoochee National Forest of

Georgia. Proceedings of the Annual Conference of the Southeast-

ern Association of Fish and Wildlife Agencies (In press).

Gentry, J. B., E. P. Odum, M. Mason, V. Nabholtz, S. Marshall, and

J. T. McGinnis. 1968. The effect of altitude and forest manipulation

on relative abundance of small mammals. Journal of Mammalogy

49:539-541.

Golley, F. B. 1966. South Carolina mammals. Contributions from

the Charleston Museum, Charleston, South Carolina.

Hall, E. R. 1981. The mammals of North America. Second edition.

John Wiley & Sons, New York, New York.

Harvey, M. J., C. S. Chaney, and M. D. McGimsey. 1991. Distribution,

status, and ecology of small mammals of the Cherokee National

Forest, Tennessee (Southern Districts). Unpublished manuscript. Center

for the Management, Utilization, and Protection of Water Resources,

Tennessee Technological University, Cookeville.

Harvey, M. J., M. D. McGimsey, and C. S. Chaney. 1992. Distribution,

status, and ecology of small mammals of the Cherokee National

Forest, Tennessee (Northern Districts). Unpublished manuscript. Center

for the Management, Utilization, and Protection of Water Resources,

Tennessee Technological University, Cookeville.

20 Joshua Laerm, et al.

Johnston, D. W. 1967. Ecology and distribution of mammals at

Highlands, North Carolina. Journal of the Elisha Mitchell Scientific

Society 83:88-98.

Junge, J. A., and R. S. Hoffmann. 1981. An annotated key to the

long-tailed shrews (genus Sorex) of the United States and Canada,

with notes on middle American Sorex. Occasional Papers of the

Museum of Natural History, University of Kansas 94:1-48.

Kalko, E. K. V., and C. O. Handley, Jr. 1993. Comparative studies

of small mammal populations with transects of snap traps and

pitfall arrays in southwest Virginia. Virginia Journal of Science

44:3-18.

Kirkland, G. L., Jr. 1985. Small mammal communities in temperate

North American forests. Australian Mammalogy 8:137-144.

Kirkland, G. L., Jr. 1991. Competition and coexistence in shrews

(Insectivora: Soricidae). Pages 15-22 in The biology of the Soricidae

(J. S. Findley and T. L. Yates, editors). The Museum of Southwestern

Biology, Albuquerque, New Mexico.

Kirkland, G. L., Jr., and P. K. Sheppard. 1994. Proposed standard

protocol for pitfall sampling of small mammal communities. Pages

277-281 in Advances in the biology of shrews (J. F. Merritt, G.

L. Kirkland, Jr., and R. K. Rose, editors). Special Publication,

Carnegie Museum of Natural History 18:1-485.

Lee, D. S., J. B. Funderburg, and M. K. Clark. 1982. A distribu-

tional survey of North Carolina mammals. Occasional Papers of

the North Carolina Biological Survey, Raleigh.

Linzey, A. V., and D. W. Linzey. 1971. Mammals of the Great

Smoky Mountains National Park. University of Tennessee Press,

Knoxville.

Mengak, M. T., D. C. Guynn, Jr., J. K. Edwards, D. L. Sanders, and

S. M. Miller. 1987. Abundance and distribution of shrews in

western South Carolina. Brimelyana 13:63-66.

Odum, E. P. 1949. Small mammals of the Highlands (North Caro-

lina) Plateau. Journal of Mammalogy 30:179-192.

Pagels, J. F. 1987. The pygmy shrew, rock shrew and water shrew:

Virginia's rarest shrews (Mammalia: Soricidae). Virginia Journal

of Science 38:364-368.

Pagels, J. F. 1991. A high elevation record for the least shrew,

Cryptotis parva (Say). Virginia Journal of Science 42:361-362.

Pagels, J. F., and C. O. Handley, Jr. 1989. Distribution of the

southeastern shrew, Sorex longirostris Bachman, in western Vir-

ginia. Brimleyana 15:123-131.

Pagels, J. F., and C. M. Tate. 1976. Shrews (Insectivora: Soricidae)

of the Paddy Knob-Little Back Creek Area of western Virginia.

Virginia Journal of Science 27:202-203.

Masked Shrew and Red-backed Vole 21

Pagels, J. F., K. L. Uthus, and H. E. Duval. 1994. The masked

shrew, Sorex cinereus, in a relictual habitat of the southern Ap-

palachians. Pages 103-109 in Advances in the biology of shrews

(J. F. Merritt, G. L. Kirkland, Jr., and R. K. Rose, editors).

Special publication, Carnegie Museum of Natural History 18:1—

485.

Pivorun, E. B., D. H. Allen, and D. T. Sawyer. 1984. First record

of Clethrionomys gapperi (Mammalia: Rodentia) in South Caro-

lina. Journal of the Elisha Mitchell Society 100:33.

Smith, C. R., J. Giles, and M. E. Richmond. 1976. The mammals

of northeastern Tennessee. Journal of the Tennessee Academy of

Science 49:88-94.

van Zyll de Jong, C. G., and G. L. Kirkland, Jr. 1989. A morpho-

metric analysis of the Sorex cinereus group in central and east-

ern North America. Journal of Mammalogy 70:110-122.

Wharton, C. H. 1968. First records of Microsorex hoyi and Sorex

cinereus from Georgia. Journal of Mammalogy 49:158.

Wharton, C. H. 1977. The natural environments of Georgia. Georgia

Department of Natural Resources, Office of Planning and Re-

sources, Atlanta.

Whitaker, J. O., Jr., G. S. Jones, and D. D. Pascal. 1975. Notes on

mammals of the Fires Creek Area, Nantahala Mountains, North

Carolina, including their parasites. Journal of the Elisha Mitchell

Scientific Society 91:13-17.

Received 25 July 1994

Accepted 30 November 1994

Rediscovery of the Aquatic Gastropod

Helisoma eucosmium (Bartsch, 1908),

(Basommatophora: Planorbidae)

William F. Adams

Environmental Resources Branch

United States Army Corps of Engineers

P.O. Box 1890, Wilmington, North Carolina 28402

AND

Susan G. Brady

Department of Biology

University of North Carolina at Wilmington

601 South College Road, Wilmington, North Carolina 28403

ABSTRACT — A population of Helisoma eucosmium (Bartsch, 1908),

a small freshwater planorbid snail considered to be extinct by

some authors, has been discovered in Town Creek, a tidal

swamp stream in southeastern North Carolina. This rediscovery

will permit a definitive determination of the proper systematic

placement of the taxon. In the absence of live specimens, past

analysis of this taxon during systematic revisions of the Planorbidae

relied only on shell morphology of type material. Under this

circumstance, this taxon was variously placed in both tropical

and temperate genera and, compounding that problem, was treated

as a full species by some authors and a subspecies by others.

Its apparently limited range elevates concern for the conserva-

tion of planorbid snail diversity in southeastern North Carolina

because it is the second taxon with a severely restricted dis-

tribution to be found in this rapidly urbanizing region.

Helisoma eucosmium (Bartsch, 1908) is a small, distinctive planorbid

snail which was collected and described from Greenfield Lake, a

millpond constructed prior to 1750 (Adams 1990a). This site is located

within the City of Wilmington, New Hanover County, North Carolina,

and is on a tributary to the lower Cape Fear River. During the past

few decades, repeated attempts to recollect H. eucosmium in Greenfield

Lake and elsewhere within the region have been unsuccessful (Fuller

1977, Adams 1990a). This fact, combined with the water quality degradation

in many regional streams, has led many investigators to treat the

taxon as extinct (Opler 1976, Imlay 1977, Palmer 1985) or possibly

extinct (Fuller 1977, Adams 19906).

Brimleyana 22:23-29, June 1995 23

24 William F. Adams and Susan G. Brady

During the spring of 1994, while performing a survey of the

mollusks of Town Creek, a tidal swamp stream tributary to the lower

Cape Fear River in adjacent Brunswick County, SGB discovered a

population of Helisoma eucosmium approximately 14.5 km (9 mi)

SSW of the type locality (Fig. 1). Although much of this stream

system remains to be investigated, the taxon is known to occupy at

least a 1.75 km (1.1 mi) stretch of the main creek. Five specimens

were deposited in the invertebrate research collection at the North

Carolina State Museum of Natural Sciences, Raleigh (NCSM). All

conform precisely to the original shell description given by Bartsch

(1908), particularly in displaying the chestnut-colored bands (Figs. 2

and 3). The largest specimen (NCSM #P1207), collected on 30 April

1994, has a greater diameter of 6.0 mm, a lesser diameter of 4.75

mm, and a height of 3.0 mm. It was probably an adult that overwintered,

a likelihood suggested by its size and the minor pitting and corrosion

of the shell surface. Three specimens taken in early July 1994 (NCSM

#P1208) are smaller; average dimensions of three specimens are maximum

diameter 3.8 mm, minimum diameter 3.0 mm, and height 1.9 mm

(n = 3). Similarly-sized specimens taken from the same period laid

eggs in captivity indicating that sexual maturity had been attained.

The molluscan community in the freshwater part of Town Creek

is diverse; species encountered to date are listed in Table 1. Continuing

surveys may disclose additional species and most of those historically

documented (Adams 1990a) from the type locality of Helisoma eucosmium

may ultimately be found.

In his description, Bartsch (1908) assigned Helisoma eucosmium

and a subspecies from Louisiana, H. eucosmium vaughani, to the

genus Planorbis Miiller 1774. F. C. Baker (1931) restricted that genus

to species of European origin and, without examination of soft tissue

anatomy, provisionally placed eucosmium in the genus Helisoma Swainson

1840. In his subsequent monograph on the Planorbidae, Baker (1945)

retained this assignment, placing it alone with Helisoma anceps (Menke,

1830) in the subgenus Helisoma s.s.; however, he made no reference

to anatomical examinations supporting this placement. On the strength

of unpublished observations by J. P. E. Morrison of live individuals

of H. e. vaughani from Louisiana, Fuller (1977) tentatively assigned

the species to the Central American genus Taphius H. & A. Adams

1855. Taphius and other planorbid genera of tropical affinity (Afroplanorbis

Thiele 1931, Biomphalaria Preston 1910, Australorbis Pilsbry 1934,

Tropicorbis Pilsbry and Brown 1914, Planorbina Haldeman 1842,

Armigerus Clessin 1884, and Platytaphius Pilsbry 1924) have long

been the subjects of taxonomic debate and divergent systematic treatment,

Helisoma eucosmium

25

Table 1. Freshwater mollusks of upper Town Creek, Brunswick County,

North Carolina, 1994.

Gastropods

Bivalves

Viviparidae

Campeloma decision (Say)

Hydrobiidae

Amnicola limosus (Say)

Gillia altilis (Lea)

Physidae

Physella hendersoni (Clench)

Planorbidae

Micromenetus dilitatus (Gould)

Helisoma eucosmium (Bartsch)

Planorbella trivolvis (Say)

Ancylidae

Laevapex fuscus (C. B. Adams)

Unionidae

Pyganodon cataracta (Say)

Villosa delumbis (Conrad)

Ligumia nasuta (Say)

Unknown taxa of the

Elliptio complanata and

Elliptio icterina complexes

Corbiculidae

Corbicula fluminea (Muller)

Sphaeriidae

Eupera cubensis (Prime)

Musculium securis (Prime)

Unidentified sphaeriids

a problem generated and sustained by years of inconclusive taxonomic

studies resulting from the great similarity in chonchology and anatomy

existing within a genus complex of worldwide distribution (Hubendick

1955, H. B. Baker 1960). While taxonomic and systematic problems

are common in the freshwater gastropods, the need for nomenclatural

stability within the tropical planorbids is a matter of great medical

and economic importance because these snails are intermediate hosts

of the human parasite Schistosoma mansoni.

In response to a petition by Wright (1962), the International

Commission on Zoological Nomenclature (1965) issued Opinion 735,

ruling that Biomphalaria is to be given precedence over the generic

names Taphius, Planorbina, and Armigera when any or all of these

names are considered to apply to the same genus. Under this ruling,

Fuller's (1977) binomen would be valid only if Taphius is determined

to be separable from Biomphalaria at the genus level. Bypassing this

issue, Burch (1982, 1989) stated Helisoma eucosmium may be a "form

or juvenile" of Helisoma anceps (Menke, 1930) and, therefore, did

not grant it specific status. Based on shell morphology and the reproductive

maturity of the material collected from Town Creek, we find no evidence

to support this possibility. Although the very oblique aperture of H.

eucosmium is suggestive of the Biomphalaria group, the lack of vertically

depressed whorls is not; therefore, we provisionally accept Baker's

26

William F. Adams and Susan G. Brady

WILMINGTON

Fig. 1. Town Creek and vicinity (* = known range of Helisoma eucosmium,

♦ = known sites of Planorbella magnifica.

(1945) designation pending definitive studies of soft tissue anatomy.

Over much of its length, Town Creek is a tidal system approximately

30-50 m wide with maximum depths varying between 5-7 m. Whereas

the lower reaches are brackish, no salinity has been detected in the

area where Helisoma eucosmium occurs. Physical parameters of the

water are pH 5.1-7.0 (* = 6.1, n = 3), conductivity 217 //mhos/cm

(n = 2), and calcium concentration 107.9 ppm (n = 2). All specimens

of H. eucosmium came from a littoral community, occurring in less

than 3 m of water, consisting of dense continuous mats of Brazilian

elodea (Egeria densa Planchon), fanwort {Cabomba caroliniana Gray),

fragrant waterlily (Nymphaea odorata Aiton), spatterdock (Nuphar

luteum (Linnaeus)), and floating-heart (Nymphoides aquatica (Walter)).

Helisoma euc osmium

27

Fig. 2. Dorsal, ventral, and apertural

views of Helisoma eucosmium

(NCSM #P1208). Lip aperture is

incomplete.

Fig. 3. Foraging Helisoma eucosmium.

28 William F. Adams and Susan G. Brady

Streambank forest cover, typical of southern swamps, consists of bald

cypress (Taxodium distichum (Linnaeus)), tupelo gum (Nyssa aquatica

Linnaeus), red maple {Acer rubrum Linnaeus), and water ash (Fraxinus

caroliniana Miller).

Our report of Helisoma eucosmium constitutes the second recent

recollection of a likely "extinct" planorbid snail in the Cape Fear

River drainage of southeastern North Carolina, the other being the

rediscovery of Planorbella magnifica (Pilsbry) in Orton Pond (Adams

1988) and in Sandhill Creek Pond (Adams 1993), approximately 14.5

km (9.0 mi) SE and 12 km (7.5 mi) ESE, respectively (Fig. 1). Because

of the heavy residential and industrial development that has occurred

in southeastern North Carolina within the past half-century, these

waterbodies might hold some of the few remaining populations of the

original freshwater molluscan fauna of the lower Cape Fear Basin. If

H. eucosmium is restricted to Town Creek and systematic research

determines that it warrants full species rank, protection under state

and federal conservation laws might be warranted. Although the Town

Creek watershed is still very rural, covered primarily in crop and

forest lands, the surrounding region is urbanizing rapidly, and the

stream will surely be impacted by this trend. Consequently, additional

surveys to determine the distribution of H. eucosmium and resolution

of outstanding systematic issues are urgently needed.

ACKNOWLEDGMENTS— Partial funding for this work has been

provided by the University of North Carolina Center for Marine Science

Research and the Wilmington District, United States Army Corps of

Engineers. We would like to thank Dr. Eric Bolen for his administrative

support, Mr. Shaun Cain for field assistance, Dr. Courtney Hackney

for his ideas and encouragement, and Dr. Rowland Shelley for his

constructive review of this manuscript. Photographs were taken by

Mr. David M. DuMond.

LITERATURE CITED

Adams, W. F. 1988. Rediscovery of Planorbella magnifica (Pilsbry)

in southeastern North Carolina. Nautilus 102(3): 125-126.

Adams, W. F. 1990a. Recent changes in the freshwater molluscan

fauna of the Greenfield Lake basin, North Carolina. Brimleyana

16:103-117.

Adams, W. F. 1990fc. Helisoma eucosmium (Bartsch, 1908). Pages

16-17 in A report on the conservation status of North Carolina's

freshwater and terrestrial molluscan fauna (W. F. Adams, editor).

North Carolina Wildlife Resources Commission, Raleigh.

Helisoma eucosmium 29

Adams, W. F. 1993. A status survey for the magnificent ramshorn

(Planorbella magnified), a freshwater snail endemic to North Carolina.

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Baker, F. C. 1945. The molluscan family Planorbidae. University of

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America. Environmental Protection Agency, EPA-600/3-82-026.

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Fuller, S. L. H. 1977. Freshwater and terrestrial mollusks. Pages

143-194 in Endangered and threatened plants and animals of North

Carolina (J. E. Cooper, S. S. Robinson, and J. B. Funderburg,

editors). North Carolina State Museum of Natural History, Ra-

leigh.

Hubendick, B. 1955. Phylogeny in the Planorbidae. Transactions of

the Zoological Society of London 28(6):453-542.

Imlay, M. J. 1977. Competing for survival. Water Spectrum 9:7-14.

International Commission on Zoological Nomenclature. 1965. Opinion

735. Bulletin of Zoological Nomenclature 22(2):94-99.

Opler, P. A. 1976. The parade of passing species: a survey of

extinctions in the U.S. Science Teacher 43(9):30-34.

Palmer, S. 1985. Some extinct molluscs of the U.S.A. Atala 13(1): 1-7.

Wright, C. A. 1962. Planorbina Haldeman, 1852, Taphius Adams

and Adams, 1955 and Armigerus Clessin, 1884 (Mollusca, Gastropoda):

Proposed Suppression under the Plenary Powers. Z.N.(S.) 1392.

Bulletin of Zoological Nomenclature 19(1):39-41.

Received 27 September 1994

Accepted 19 January 1995

Effects of a Clearcut on the Herpetofauna of a

South Carolina Bottomland Swamp

Joseph P. Phelps1

North Carolina State University

Hardwood Research Cooperative

Raleigh, North Carolina 27695-8008

AND

Richard A. Lancia

Department of Forestry

North Carolina State University

Raleigh, North Carolina 27695-8002

ABSTRACT — Amphibians and reptiles were trapped during summer

in a South Carolina bottomland swamp to assess the impacts

of clearcut timber harvesting. Animals were captured using drift

fences with pitfall traps, coverboards, and polyvinyl chloride

pipes which simulated treefrog habitat. Twenty-nine species (10

amphibians and 19 reptiles) were detected on the site. Some

were captured frequently enough to infer microhabitat preferences.

Salamanders were much more frequent in the control area than

in the clearcut. Other species showing preferences for the control

were bronze frogs (Rana clamitans), gray treefrogs (Hyla chrysoscelis),

and box turtles (Terrapene Carolina). Reptiles generally preferred

the clearcut. This was especially true of lizards and large snakes.

Diversities showed no significant differences between the control

and clearcut. Small clearcuts done on long rotations are recommended.

Machinery impact should be kept to a minimum, and down

wood and snags should be left on the site.

Bottomland hardwood forests have been recognized for their

importance in floodwater and sediment retention, water quality protection,

timber production, and wildlife habitat (Brinson et al. 1981, Clark

and Benforado 1981, Harris and Gosselink 1986). At the same time,

these ecosystems are being lost and degraded rapidly (Turner et al.

1981, Rudis 1993) due in large part to fragmentation. In the South-

east, 75,000 acres of forested wetlands have been lost since 1982

(Cubbage and Flather 1993), not including acreage that was logged

and regenerated. These logged wetlands may temporarily lose some

functional value, and they contribute to fragmentation.

1 Present address: Weyerhaeuser Company, Southern Forest Research, P.O. Box 1060,

Hot Springs, Arkansas 71902.

Brimleyana 22:31-45, June 1995 31

32 J. P. Phelps and R. A. Lancia

The research results presented here were obtained concurrent

with other studies (Pavel 1993, Perison In Press) designed to document

the impact of timber harvesting on the functional value of a bottom-

land swamp. Amphibians and reptiles (herpetofauna) were chosen as

the appropriate wildlife groups to study because of their abundance in

the Southeast (Keister 1971, Vickers et al. 1985, Hairston 1987) and

because of their importance in food chains (Pough et al. 1987, Blaustein

and Wake 1990). Herpetofaunal species are also influenced by factors

that are affected by timber harvest, including hydrology, soil quality,

and vegetative structure. Herpetofaunal communities have been shown

to be altered by clearcutting (Enge and Marion 1986, Pough et al.

1987, Petranka et al. 1993), ditching of wetlands (Vickers et al. 1985,

Enge and Marion 1986), and changing forest cover (Bennett et al.

1980, Pough et al. 1987).

In addition, much attention has been paid to a possible world-

wide decline in amphibian diversity (Blaustein and Wake 1990, Pechmann

et al. 1991, Hairston and Wiley 1993). Logging has been identified

among the many possible causes of such a decline (Wake 1991, Hairston

and Wiley 1993). Amphibians may be a good indicator of general

environmental degradation, due to their exposure to terrestrial and

aquatic toxins, and their sensitivity to habitat changes (Beiswenger

1988, Blaustein and Wake 1990).

The objectives of our research were to evaluate whether clear-

cutting in a hardwood swamp had any effect on community diversity

or abundance of summer-active amphibians and reptiles. We also attempted

to identify habitat variables that may have been related to changes in

the herpetofaunal community.

METHODS

Study Site

The study site was on the South Fork of the Edisto River,

near Norway (Orangeburg County), South Carolina. "The site is re-

presentative of blackwater swamps in the Carolinas that have timber

management potential" (Perison et al. 1993). Predominant trees in the

swamp forest included tupelo gum (Nyssa spp.), sweetgum (Liquidambar

styraciflua), willow/water oak (Quercus phellos/nigra), and green ash

(Fraxinus pennsylvanica) (Pavel 1993). A clearcut of approximately

10 ha was completed in January 1991. Much of the clearcut area was

impacted by skidder tire ruts. The adjacent control area was upstream

of the clearcut and was a second growth stand approximately 45 years

old. Second growth forest surrounded the clearcut. The sampled area

of control was comparable to the size of the clearcut. The edge sampled

was 650 m long.

Clearcut Herpetofauna 33

Herpetofauna

Sampling — Amphibians and reptiles were captured using three

types of traps: drift fences, coverboards, and polyvinyl chloride (PVC)

pipe. Nine arrays of drift fences, measuring 270 m in total, were

constructed of aluminum flashing (Gibbons and Semlitsch 1981).

Each array consisted of two 15-m lengths placed at right angles to

one another. Three of the arrays were placed in the clearcut, three in

the control, and three along the edge between the clearcut and control.

Twelve pitfall traps (19-L plastic paint buckets) were placed along

each array, for a total of 108 traps. Arrays were centered in the

clearcut, about 75 m from the edge. Control arrays averaged about

75 m from the edge.

Forty-five coverboards were systematically placed across the study

site, 30 in the clearcut and 15 in the control. Each coverboard con-

sisted of a piece of plywood or particle-board (about 120 x 60 x 0.625

cm) placed flat on the ground to simulate the type of cover often used

by ground-dwelling herpetofauna (DeGraff and Yamasaki 1992, Fitch

1992, Mitchell et al. 1993). Coverboards were placed a minimum of

20 m apart.

Sixty 1.5-m lengths of PVC pipe were driven into the soil to

capture treefrogs. The diameters of the pipes were 2.5 and 5 cm.

Forty pipes were used in the clearcut, and 20 in the control. These

pipes served as refuges for treefrogs, which were easily captured at

the open top ends of the pipes. One nocturnal chorus survey (July

1993) was done to compare to habitat use trends indicated by the pipe

captures. Frog choruses were monitored from the clearcut, control,

and edge for 30 minutes each. Weather was humid, warm, clear, and

calm.

Each of the traps was checked daily during the summers of 1992

and 1993. All captured animals were released immediately. Adult

anurans were toe-clipped, but no capture-recapture data will be pre-

sented here (see below). In 1992, we trapped from 29 May until 13

August, with the exception of 14 days when the swamp was flooded.

On those days, only the treefrog pipes were checked. In 1993, four

drift fence arrays (one control, two edge, one clearcut) were checked

from 20 May to 12 August. The remaining traps were checked from

20 June to 12 August. Again, sampling was impeded due to flooding.

Sampling was replicated within the control, edge, and clearcut

site, but not replicated with additional sites because of constraints

imposed by the establishment of concurrent studies, which used small

replicated treatment blocks within the clearcut (Pavel 1993, Phelps

1993, Perison In Press). These blocks were the reason for unequal

34 J. P. Phelps and R. A. Lancia

sampling effort for coverboards and PVC pipes in the clearcut, edge,

and control. Due to the lack of replication, the results and conclusions

from this study apply to the particular site we studied. Generalization

to other sites is risky.

Data analysis — Capture data were analyzed using the Shannon-

Weaver Diversity Index (Poole 1974). This allowed statistical com-

parisons among drift fence arrays in the clearcut, on the edge, and in

the control; and between coverboards in the clearcut and in the control.

Variance for each treatment was calculated based on the number of

captures in each treatment, and was used to calculate the ^-statistic (a

= 0.05) for each comparison (Poole 1974). The formulas used are as

follows:

Var (//') = [rMPl ln2p,] - [g.lf>, lnp,]2 + s-1

N 2N2

H\-H\

[Var(/f ,) + Var(//'2)]1/2

d.f. = [Var(/f ,) + Var(/r2)]2

[VarCtf',)]2 + [Var(tf'2)]:

N, N2

where i represents each species in the sample, pi = proportion of

species i in the sample, s = number of species in the sample, N =

number of captures in the sample.

Assumptions of the Shannon-Weaver Index were that all species

present were sampled, and all were equally catchable. These assumptions

were not met because of the differing ability of the traps to catch

various species (Gibbons and Semlitsch 1981). Also, two species caught

by hand were never captured in a trap of any kind and were therefore

excluded from calculations of H' . These were the brown water snake

(Nerodia taxispilota) and the timber rattlesnake (Crotalus horridus).

Their size explains the lack of pitfall captures, as no larger snakes

were caught in pitfalls. Other large snakes were captured under

coverboards.

Poole (1974) states that "no great error" will occur in calculating

Shannon-Weaver as if all the species available are present in the sample,

even when, as in this case, some species are not represented. Unequal

catchability of species should still allow relative diversity comparisons.

Clearcut Herpetofauna 35

The lack of replication allowed only a rough comparison of

capture frequencies, not a statistical test of differences in abundance

(Hurlbert 1984). The original intent was to estimate abundances of

anurans using mark-recapture methods, but a lack of sufficient recapture

prevented this. To make direct comparisons of capture frequencies,

we assumed that capture probabilities for each species were the same

across habitats. This assumption could not be tested, so habitat

preferences should be interpreted with caution.

Habitat Characteristics

Sampling — In an attempt to relate herpetofaunal diversity to

microhabitat variability, the percent cover of midstory and overstory

trees was measured with a spherical densiometer at five points around

each drift fence array. One reading was taken in the center and one at

a distance of 15 m in each cardinal direction. Percent cover of veg-

etation less than 2 m tall was also measured, using a line intercept

method (Barbour et al. 1987). There were four 15-m transects at each

drift fence array, originating at the middle of the array and extending

in each cardinal direction. A complete description of the vegetation

of the site is given by Pavel (1993).

The cross-sectional area of coarse woody debris was measured

using the same transects. In this case, the transects were thought of

as vertical planes that extended from the ground to the highest piece

of downed wood. Each piece of wood greater than 8-cm diameter (at

the point of intersection) was measured to the nearest centimeter and

classified as "sound" or "rotten" (Brown 1974).

Surface soil temperatures and soil densities were measured by

Perison (In Press) in the control and clearcut areas. All habitat data

were analyzed using Mests at the 0.05 alpha level. Each measurement

was considered to be independent, and means and standard errors

were calculated for each variable. In the clearcut, soil temperature

and density were measured in areas with skidder traffic, but not in

skidder ruts. These variables were not measured at the edge.

RESULTS

Herpetofauna

Shannon-Weaver Diversity Indices calculated for the drift fence

and coverboard captures are given in Table 1. There was no significant

difference between clearcut and control diversity indices for either of

the two trapping methods. The diversity of the edge drift fences was

lower than either the control (t = 8.70, d.f. = 1,538) or clearcut drift

fences (t = 9.70, d.f. = 2,209). The reason for this difference was the

36

J. P. Phelps and R. A. Lancia

Table 1. Summary of herpetofauna capture data, Edisto River swamp,

South Carolina, 1992-1993. H' is the value of the Shannon-Weaver

diversity index. Values of H' with the same letter were not different

at 5% alpha level.

Captures Species

(N) (s)

H'

Control drift fences

Clearcut drift fences

Edge drift fences

All drift fences

Control coverboards

Clearcut coverboards

All coverboards

heavy weighting toward one species, the southern toad {Bufo terrestris),

on the edge. Clearcut and control drift fence diversities were not

different at the 0.05 alpha level. Coverboard diversities were higher

than drift fence diversities (f = 18.5, d.f. = 333). Drift fences captured

more species (23) and individuals (4,749) than coverboards (19 and

279, respectively). The inclusion of the clearcut increased the richness

of the capture sample. Twenty-three species were captured on the site

in drift fences, but only 16 species were captured in the clearcut, and

16 in the control. Nineteen species were captured under coverboards,

but only 16 were captured in the clearcut, and nine in the control.

Several species showed clear preferences for either the control

or the clearcut (Table 2). All types of salamanders were detected

more often in the control area. A total of 112 salamanders was captured

in pitfall traps and under coverboards. Ninety-two (82.1%) of these

were captured in the control area, with only 9 (8.0%) in the clearcut

(Table 2). Bronze frogs {Rana clamitans) were captured more frequently

in the control (104 control captures to 17 in the clearcut), while

southern leopard frogs (R. utricularia) were less common in the control

(33 drift fence captures compared to 52 on the edge and 51 in the

clearcut). Other frog species showing preferences were eastern narrow-

mouth toads {Gastrophryne carolinensis) and green treefrogs (Hyla

cinerea), which preferred the clearcut, and gray treefrogs (H. chrysoscelis),

which preferred the control. Southern toads, most of which were

juveniles, were captured most frequently in the edge pitfalls. Southern

toads were abundant in all three areas. The data on frog and toad

species were supported by the breeding chorus monitoring.

Clearcut Herpetofauna 37

Reptiles generally seemed to prefer the clearcut. Eastern mud

turtles (Kinosternon subrubrum) were captured 41 times in the clearcut,

35 times on the edge, and only 10 times in the control area. Common

musk turtles {Sternotherus odoratus) and eastern box turtles (Terrapene

Carolina) showed the opposite trend. Lizards (Eumeces fasciatus and

Anolis carolinensis) and large snakes (chiefly Agkistrodon piscivorus

and Nerodia spp.) were more common in the clearcut. Because they

were rarely captured in traps but often seen, habitat preferences of

large snakes, lizards, and box turtles are largely inferred from hand

captures, rather than trapping data.

Habitat Characteristics

Microhabitat variables measured in each area are given in Table

3. Over- and mid-story canopy cover was highest in the control (95%),

followed by the edge (74%), and clearcut (6%). Understory canopy

cover followed the reverse trend, being highest in the clearcut (95%),

followed by the edge (48%), and the control (5%).

Cross-sectional area of sound coarse woody debris was significantly

higher in the clearcut, and not significantly different between the

edge and control. The mount of rotten coarse woody debris was not

different among the three areas. Soil surface temperature and soil

compaction were not measured on the edge, but were not significantly

different between the clearcut and the control, or between rutted and

non-rutted areas in the clearcut.

DISCUSSION

Habitat Preferences

Fewer salamanders were captured in the clearcut, as compared

to the control. High temperatures and insolation, and low relative

humidity may all have contributed to this, as these factors increase

the risk of desiccation. Salamanders need to keep their skin moist for

gas exchange, and their large surface areas to volume ratios make

moisture retention difficult (Duellman and Trueb 1986). Moisture is a

key factor in determining where salamanders can live (Wyman 1988,

Petranka et al. 1993). Petranka et al. (1993) speculated that 75% to

80% of southern Appalachian salamanders die of physiological stress

due to desiccation following clearcuts.

Ecological differences may explain the varying preferences of

frog species. Southern toads are generalists, and their common

occurrence along the edge suggests that individuals were using both

habitats. Bronze frogs and southern leopard frogs showed opposing

preferences, for the control and clearcut respectively. Bronze frogs

38

J. P. Phelps and R. A. Lancia

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are a forest wetland species, whereas leopard frogs are more ubiquitous

(Mount 1975). Enge and Marion (1986) found fewer southern leopard

frogs in clearcuts than in control areas in southern pine flatwoods.

Eastern narrowmouth toads emerge from burrows only when ground

water is sufficiently pooled for breeding. Reduced evapotranspiration

in the clearcut (Perison et al. 1993) may have increased the likelihood

of these frogs receiving the cue to emerge. Skidder tire ruts may have

had a similar effect on these and the other frog species, as they were

a source of fish-free breeding pools (Phelps 1993).

Two species of treefrogs were captured, and these showed opposing

preferences. Green treefrogs may have predominated in the clearcut

because the traps (treefrog pipes) were level with their habitat (leafy

sprouting vegetation). Green treefrogs may have been equally abundant

in the control (as suggested by the number of control captures, Table

2), but may have been living higher than the traps were sampling.

Gray treefrogs were rarely captured, and the reason again may have

been that the traps were too low. The removal of trees in the clearcut

may have excluded gray treefrogs because their habitat (tree boles)

was removed. In the case of the green treefrogs in the clearcut, their

habitat quickly returned, but at a lower height. Overall, the treefrog

pipe method showed promise, as it was inexpensive and trapped

significant numbers of frogs while causing them no apparent stress.

Reptiles may have preferred the clearcut because of increased

temperature and insolation. A basking reptile would be at an advantage

in the direct sun of the clearcut because it could achieve active temperatures

faster and remain active longer through the day. Bury and Corn (1988),

working in the Pacific Northwest, found that "reptiles predominate in

clearcuts, most likely responding to increased ambient tempera-

tures in such areas." The presence of a large amount of slash in the

clearcut may have provided a valuable habitat component in the form

of cover for reptiles or their prey. Reptiles that seemed to prefer the

control included ringneck snakes (Diadophis punctatus), copperheads

(Agkistrodon contortix), common musk turtles, and eastern box turtles.

The reasons for these preferences are not clear, although ringneck

snakes were the smallest and most fossorial reptile species regularly

captured, and may have been less dependent on basking and more

dependent on an undisturbed forest floor.

Recommendations and Conclusions

Clearcut size is one factor to be considered when planning for

natural forest regeneration in forested wetlands. Amphibians and reptiles

have relatively small home range sizes (Duellman and Trueb 1986)

and, therefore, cannot disperse from or quickly recolonize impacted

Clearcut Herpetofauna 41

areas. Hairston (1987) found that salamanders return to their home

range even after disturbance and handling. Small clearcuts with undisturbed

sources of recolonization nearby were advocated by Buhlmann et al.

(1988) and Enge and Marion (1986). A mosaic of small clearcuts,

second growth, and undisturbed areas would likely create increased

landscape diversity, as compared to a single homogeneous stand.

Recovery times of about 60 years for salamander populations in

clearcut areas are given by Petranka et al. (1993) in the southern

Appalachian and Pough et al. (1987) in New York. This suggests that

long rotation times are needed to avoid a long-term decline of sala-

mander populations over several rotations. Enge and Marion (1986)

also recommended long rotation times that allow adequate recovery

of herpetofaunal populations.

Controlling certain aspects of the harvest operation can minimize

the adverse effects of a clearcut. Most importantly, snags and coarse

woody debris should be left on the site (Enge and Marion 1986).

Woody debris should be of large size and in an advanced state of

decay (Bury and Corn 1988, Welsh and Lind 1988). Aubry et al.

(1988) suggested that "the abundance levels of salamanders are more

likely a function of the availability of woody debris for cover than

age of the overstory." Since suitable woody debris is more abundant

in older stands, longer rotation times are important. Leaf litter on the

forest floor is another important component of herpetofaunal habitat

(Pough et al. 1987, DeGraaf and Rudis 1990, Petranka et al. 1993),

and can be destroyed by ground machinery such as skidders (Buhlmann

et al. 1988). Skidders should be restricted to small areas, and helicopters

should be used when practical. Buhlmann et al. (1988) recommended

harvesting in the season of inactivity for the local herpetofauna, but

some southern Coastal Plain species are active at all times of the

year.

Further studies in which capture probabilities can be estimated

would allow direct comparison of capture data among species in the

same habitat, and within species across habitats. Remaining research

opportunities include determining the fate of salamanders in the face

of clearcutting, and monitoring subsequent recovery of populations

through recolonization. Also, a study similar to ours, focusing on

winter-active amphibians, would be valuable. In addition, the habitat

value of skidder ruts should be studied. The possible benefit of extra

standing water (Phelps 1993) may mitigate the effects of soil degradation

(Buhlmann et al. 1988). Finally, additional work with PVC pipes for

capturing treefrogs should be done, including their use in various

habitats, with different species, and the possible effect of pipe height

on trap efficiency.

42 J. P. Phelps and R. A. Lancia

The key to expanding knowledge in the area of wildlife habitat/

forestry relationships is to replicate treatments. In this case, conclusions

could have been strengthened by having several clearcuts and several

control areas (Hurlbert 1984). Specific factors such as woody debris,

size of clearcuts, and skidder rut impact could be studied with such a

design. Ideally, each area would be sampled prior to the installation

of the clearcut (Buhlmann et al. 1988). This would allow comparison

of data from the clearcut before and after treatment, and from the

control. Effects of space, time, and the treatment itself could be separated.

Replication and the collection of baseline data could be achieved

more easily within the framework of Adaptive Resource Management

(Walters 1986). This is a system of research integrated with management,

wherein management decisions are treated as hypotheses and tested

with replicated trials. After several iterations of hypothesis and experiment,

predictions involved with policy can become prescriptions based on

hard data.

ACKNOWLEDGMENTS— funding for this project was provided

by the Hardwood Research Cooperative of North Carolina State University.

Harvest treatments were done by Georgia-Pacific Corporation, and

housing for researchers was provided by the South Carolina Forestry

Commission. Dr. Robert Kellison of the Hardwood Cooperative provided

valuable support and a review of the manuscript. Two anonymous

reviewers provided valuable comments.

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Clark, J. R., and J. Benforado (editors). 1981. Wetlands of bottom-

land hardwood forests. Elsevier Publishing Company, Amsterdam,

Netherlands.

Cubbage, F. W., and C. H. Flather. 1993. Distribution of and trends

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seventh biennial southern silvicultural research conference (J. C.

Brissett, editor). United States Department of Agriculture Forest

Service, General Technical Report SO-93.

DeGraaf, R. M., and D. D. Rudis. 1990. Herpetofaunal species com-

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Duellman, W. E., and L. Trueb. 1986. Biology of amphibians. McGraw-

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Enge, K. M., and W. R. Marion. 1986. Effects of clearcutting and

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Forest Ecology and Management 14:177-192.

Fitch, H. S. 1992. Methods of sampling snake populations and their

relative success. Herpetological Review 23:17-19.

44 J. P. Phelps and R. A. Lancia

Gibbons, J. W., and R. D. Semlitsch. 1981. Terrestrial drift fences

with pitfall traps: an effective technique for quantitative sampling

of animal populations. Brimleyana 7:1-16.

Hairston, N. G. 1987. Community ecology and salamander guilds.

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Hairston, N. G., and R. H. Wiley. 1993. No decline in salamander

(Amphibia: Caudata) populations: a twenty year study in the southern

Appalachians. Brimleyana 18:59-64.

Harris, L., and J. G. Gosselink. 1986. Cumulative impacts of bot-

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terrestrial wildlife. United States Environmental Protection Agency.

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lina State University, Raleigh.

Pechmann, J. H. K., D. E. Scott, R. D. Semlitsch, J. P. Caldwell, L.

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Clearcut Herpetofauna 45

Poole, R. W. 1974. An introduction to quantitative ecology. McGraw-

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1988:394-399.

Received 16 November 1994

Accepted 19 January 1995

First Record of the Water Shrew, Sorex palustris

Richardson (Insectivora: Soricidae), in Georgia

with Comments on its Distribution and Status

in the Southern Appalachians

Joshua Laerm, Charles H. Wharton

Museum of Natural History and Institute of Ecology

AND

William Mark Ford

Daniel B. Warnell School of Forest Resources

University of Georgia, Athens, Georgia 30602

ABSTRACT— The first state record of Sorex palustris is

reported from Georgia, in a markedly boreal habitat in the

upper headwaters of the Tallulah River in Towns County. Records

in the southern Appalachians indicate the species to be rare

and its distribution characterized by a series of disjunct

populations.

On 30 May 1994 one adult male water shrew (Sorex palustris

Richardson) was recovered from a sunken pitfall trap adjacent to

Mare Cove Branch at its junction with Burnt Cabin Branch, a tributary

of the Tallulah River in extreme northern portion of Towns County,

Georgia, at an elevation of 808 m. The specimen was recovered under

a rotting log and other woody debris immediately adjacent to the base

of a 25 m waterfall in rocky talus. Standard body measurements were:

138-64-19. This is the first record of the species from Georgia and

represents an extension of its range approximately 25 km southeast

from its nearest reported locality along a short section of Fires Creek

in Clay County, North Carolina, at an elevation of 1,160 m (Whitaker

et al. 1975)

During the period 30 October 1993 through 30 May 1994 we

established a transect of 20 pitfall traps (a total of 5,420 trap nights)

along Burnt Cabin Branch. Pitfalls were 946 cm3 plastic cups

(11-cm lip diameter and 14-cm depth) filled with approximately

0.13-L formalin solution and set flush to the ground adjacent to

fallen logs, rocks stumps, or other forest floor debris within 5 m of

the stream edge. Traps were checked biweekly. The collection locality,

which was selectively logged in the past, is a mature, predominantly

northern hardwood forest community dominated by yellow birch

Brimleyana 22:47-51, June 1995 47

48 Joshua Laerm, C. H. Wharton, and W. M. Ford

(Betula luted), black birch (B. lenta), liden {J ilia heterophylla), and

hemlock (Tsuga canadensis), with a rhododendron (Rhododendron

maximum) understory.

The mammalian fauna of Burnt Cabin Branch has marked boreal

affinities. Wharton (1968) reported the first Georgia records of the

masked shrew (Sorex cinereus) and pygmy shrew (S. hoyi) from Beech

Creek, another tributary of the Tallulah River within 500 m of the

present locality, and Laerm (1992) reported the first Georgia record

of the hairy-tailed mole (Parascalops brewerii) from the present locality.

Other small mammals recovered in pitfalls and snap traps at Burnt

Cabin Branch include Sorex cinereus, S. fumeus, Blarina brevicauda,

Tamiasciurus hudsonicus, Peromyscus maniculatus, Napaeozapus

insignis, and Clethrionomys gapperi.

Sorex palustris is distributed in the transcontinental Canadian

boreal forest from Nova Scotia westward to southeastern Alaska and

southward throughout much of the Sierra Nevada and Rocky

Mountains in the western United States as well as the Appalachian

Mountains to Tennessee, North Carolina (Hall 1981, Beneski and

Stinson 1987), and now Georgia in the eastern United States. Populations

throughout the Appalachian Mountains from southwestern Pennsylvania

to Georgia are referable to S. palustris punctulatus Hooper 1942, the

West Virginia water shrew. Based on available published sources,

museum records, and personal communications it appears that S. p.

punctulatus is rare and its distribution characterized by a series of

apparently disjunct populations.

The northernmost record for S. p. punctulatus is a single specimen

from Cove Run in the Negro Mountains, Somerset County, Pennsyl-

vania (Doutt et al. 1966, Enders 1985). More recently, two additional

specimens have been obtained from Somerset County (C. Bier and

S. McLaren, personal communication; specimens in Carnegie Museum

of Natural History). Apparently, the distribution of this subspecies is

disjunct from that of S. p. albibarbis which is reported from central

and northeastern Pennsylvania and northward (Hall 1981, Beneski

and Stinson 1987, Merritt 1987).

Mansuetti (1958), Paradiso (1969), and Feldhamer et al. (1984)

discussed the questionable occurrence of S. palustris in Maryland.

However, seven individuals of S. p. punctulatus are now known from

seven sites in Maryland, all from Garrett County (E. Thompson,

Maryland Natural Heritage Program, personal communication). At least

12 individuals are known from five counties (Pendleton, Pocahantas,

Preston, Randolph, and Tucker) in West Virginia (Kellogg 1937,

Hooper 1942, McKeever 1952, and records on file with West Virginia

Water Shrew 49

Heritage Inventory Program). Three individuals are known from a

single locality in Bath County, Virginia, and five from three localities

in Highlands County, Virginia (Pagels and Tate 1976, Pagels 1987,

Handley 1991, and J. Pagels, personal communication). Thirteen

records are known from the Great Smoky Mountains National Park,

Sevier County, Tennessee (Conaway and Pfitzer 1952, Linzey and

Linzey 1968), and Harvey et al. (1991) report an additional 18 speci-

mens from four localities in Monroe County, Tennessee. In North

Carolina it is known from five individuals from Clay County and one

specimen from Great Smoky Mountains National Park in Swain

County (Whitaker et al. 1975, Linzey 1983, Webster 1987).

Compared to other soricids in the southeastern United States,

Sorex palustris appears to be rare. Sorex p. punctulatus is considered

a Category 2 taxon by the United States Fish and Wildlife Service, is

listed as endangered in Virginia (Virginia Department of Game and

Inland Fisheries), and is considered a species of special concern

in North Carolina (North Carolina Natural Heritage Program) and

Tennessee (Tennessee Wildlife Resource Agency). Although other

soricids, such a Sorex hoyi and Sorex dispar, that historically have

been considered extremely rare are now known to be more widely

distributed and more common than previously believed (Pagels 1987,

Handley 1991, Laerm et al 1994), the water shrew appears to be the

rarest and most localized shrew in the southeastern United States.

Additional surveys for the water shrew are required to assess its

true status.

ACKNOWLEDGMENTS— This study was supported through a

cooperative funding agreement between the United States Forest

Service, Chatahoochee National Forest, and The University of

Georgia Museum of Natural History.

LITERATURE CITED

Beneski, J. T., Jr., and D. W. Stinson. 1987. Sorex palustris. Mam-

malian Species 296:1-6.

Conaway, C. S., and D. W. Pfitzer. 1952. Sorex palustris and Sorex

dispar from the Great Smoky Mountains National Park. Journal

of Mammalogy 33:106-108.

Doutt, J. K., C. A. Heppenstall, and J. E. Guilday. 1966. Mammals

of Pennsylvania. The Pennsylvania Game Commission, Harrisburg.

50 Joshua Laerm, C. H. Wharton, and W. M. Ford

Enders, J. E. 1985. Water shrew Sorex palustris punctulatus (Hooper).

Pages 383-385 in Species of special concern in Pennsylvania (H.

H. Genoways and F. J. Brenner, editors). Special Publication of

the Carnegie Museum of Natural History, Number 11, Pittsburgh,

Pennsylvania.

Feldhamer, G. A., J. E. Gates, and J. A. Chapman. 1984. Rare,

threatened and extirpated mammals from Maryland. Pages 395-438

in Threatened and endangered plants and animals of Maryland

(A. W. Norden, D. C. Forester, and G. H. Fenwick, editors).

Maryland Natural Heritage Program Special Publication 84-1.

Hall, E. R. 1981. The mammals of North American. Second edition.

John Wiley & Sons, New York, New York.

Handley, C. O., Jr. 1991. Mammals. Pages 539-613 in Virginia's

endangered species (K. Terwilliger, editor). McDonald and Woodward,

Blacksburg, Virginia.

Harvey, M. J., C. S. Chaney, and M. D. McGimsey. 1991. Distribu-

tion, status, and ecology of small mammals of the Cherokee Na-

tional Forest, Tennessee (Southern Districts). Report to the United

States Forest Service. Manuscript on file, Center for the Manage-

ment, Utilization, and Protection of Water Resources, Tennessee

Technological University, Cookville.

Hooper, E. T. 1942. The water shrew {Sorex palustris) of the south-

ern Allegheny Mountains. Occasional Papers of the Museum Zo-

ology, University of Michigan 463:1-4.

Kellogg, R. 1937. Annotated list of West Virginia mammals. Pro-

ceedings of the United States National Museum 84:443-479.

Laerm, J. 1992. Georgia's rarest mammal. Georgia Wildlife 2(2):46-53.

Laerm, J., W. M. Ford, and D. W. Weinand. 1994. Additional

records of the pygmy shrew, Sorex hoyi winnemana Preble (In-

sectivora: Soricidae), in western North Carolina. Brimleyana 21:91-

96.

Linzey, D. W. 1983. Status and distribution of the northern water

shrew {Sorex palustris) and two subspecies of northern flying

squirrel {Glaucomys sabrinus coloratus and Glaucomys sabrinus

fuscus). Final report under United States Fish and Wildlife Ser-

vice Contract No. 14-16-005-79-068.

Linzey, D. W., and A. V. Linzey. 1968. Mammals of Great Smoky

Mountains National Park. Journal of the Elisha Mitchell Scien-

tific Society 84:384-414.

Mansueti, R. 1958. The Craneville Pine Swamp. Atlantic Naturalist

13:72-84.

McKeever, S. 1952. A survey of West Virginia mammals. West

Virginia Conservation Commission, Pitman-Roberstson Project 22-

R. Unpublished report on file, United States National Museum of

Natural History, Washington, D.C.

Water Shrew 51

Pagels, J. F. 1987. The pygmy shrew, rock shrew, and water shrew:

Virginia's rarest shrews (Mammalia: Soricidae). Virginia Journal

of Science 38:364-368.

Pagels, J. F., and C. M. Tate. 1986. Shrews (Insectivora: Soricidae)

of the Paddy Knob-Little Back Creek Area of western Virginia.

Virginia Journal of Science 27:202-203.

Paradiso, J. L. 1969. Mammals of Maryland. North American Fauna

66:1-193.

Webster, W. D. 1987. Sorex palustris punctulatus. Pages 36-38 in

Endangered, threatened and rare fauna of North Carolina. Part 1.

A reevaluation of the mammals (M. K. Clark, editor). Occasional

Papers of the North Carolina Biological Survey, Raleigh.

Wharton, C. H. 1968. First records of Microsorex hoyi and Sorex

cinereus from Georgia. Journal of Mammalogy 49:158.

Whitaker, J. O., Jr., G. S. Jones, and D. D. Pascal, Jr. 1975. Notes

on the mammals of the Fires Creek Area, Nantahala Mountains,

North Carolina, including their ectoparasites. Journal of the Elisha

Mitchell Scientific Society 91:13-17.

Received 25 July 1994

Accepted 26 January 1995

Florida Manatees, Trichechus manatus

(Sirenia: Trichechidae),

in North Carolina 1919-1994

Frank J. Schwartz

Institute of Marine Sciences

University of North Carolina

Morehead City, North Carolina 28557

ABSTRACT— Florida manatees, first reported in 1919 from North

Carolina, are now known to have frequented 59 sites (68 indi-

viduals) during the period of 1919-1994. All but two have

been subadults of about 1.8-2.4-m lengths. Only seven deaths

have been recorded. Eleven coastal counties have harbored manatees.

Four occurrences have been at inlets and six in the open ocean.

Pelletier Creek, a Carteret County tributary of Bogue Sound,

along with the Atlantic Ocean have been the most frequented

sites (6); eight manatees occurred at a lush vegetation site in

the Trent River (Craven County), a tributary of the Neuse

River. Four records came from Wrightsville Beach and Sound,

three manatees entered the state from Chesapeake Bay via the

canal and Intracoastal Waterway into Currituck Sound. Farthest

inland river penetrations have been 94.4 km, 6.4 km north of

Wilmington; 92 km, Neuse River at Fort Barnwell Bridge, 33

km northeast of New Bern (Craven County); and one each

penetrated the Tar River at Washington (58 km, Beaufort County)

and Greenville (88 km, Pitt County). The increased frequency

of occurrences in later years may be the result of an in-

creased public awareness of the federally-protected species rather

than a seemingly increasing population.

The Florida manatee, Trichechus manatus, can attain a size of

4.1 m, 1,620 kg, and ranges from Maryland (Chester River, Chesapeake

Bay) to Louisiana in the northern Gulf of Mexico (C. Beck, National

Biological Service, personal communication; Jefferson et al. 1993).

Northernmost manatee records in North Carolina have been in

Currituck Sound (Dare County) (Brimley 1905, 1946; Brimley 1931;

Caldwell and Golley 1965) (Table 1, Fig. 1). My study summarizes

early records, adds 44 new records (total 59) and comments on where,

when, and what size manatees have occurred in North Carolina.

Brimleyana 22:53-60, June 1995 53

54

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57

Fig. 1. Locations for 68 manatee sightings in North Carolina during

the years 1919-1994.

EARLY LITERATURE OBSERVATIONS

Fifteen live and three dead manatees were reported (16 sites)

from North Carolina between 1919 and 1976 by Brimley (1931, 1946),

Lee (1976), Rathbun et al. (1981), and Smith et al. (1960). The specimen

from Currituck Sound near Duck (Brimley 1946) was about 3 m long.

None was weighed, although a Wrightsville Sound specimen and

the Currituck specimens were estimated to weigh 450 kg (Brimley

1931, 1946). Clark (National Marine Fisheries Service, personal

communication) recorded a live manatee near Cape Lookout 25 June

1975.

58 Frank J. Schwartz

RECENT NORTH CAROLINA OBSERVATIONS

Manatees are now known (68 individuals from 59 sites) to fre-

quent nearly all North Carolina ocean and inland waters (Table 1,

Fig. 1). From 1977 to 1994, C. Beck (National Biological Service,

personal communication) reports three live and three dead manatees

from six sites between 1982 and 1990; Clark adds two additional live

individuals in 1982 and 1987 and the remaining 42 specimens (41

sites) are from my study (Table 1). All but four occurrences have

been of live specimens. Recent manatee sightings have been of sub-

adults or young about 1.8-2.4 m long (Table 1). None has been weighed.

Manatees have been recorded from 11 coastal North Carolina coun-

ties (Table 1); 15 occurrences have been in Carteret County, nine

each in Dare and New Hanover, and eight in Craven County. Most

sightings occurred in 1983 and 1994 (nine), five in 1976, and two to

four sightings during most other years. Sightings have occurred dur-

ing nine months of the year, mostly in September (14), followed by

eight in August and October (Table 1). Most often frequented locali-

ties have been: Pelletier Creek (six), a tributary to Bogue Sound at

Morehead City (Carteret County), various localities in the Atlantic

Ocean (6), four at Wrightsville Beach and Sound (New Hanover County),

and the Trent River near New Bern, North Carolina (8) (Craven County).

While Rathbun et al. (1981) noted that open ocean habitat occur-

rences are rare, four North Carolina records have been from inlet-

ocean sites and six in the open ocean. Of the latter, three ocean

occurrences were off Shackleford Banks (Carteret County), one off

Wrightsville Beach (New Hanover County), and one each off the

Outer Banks off Avon and Kitty Hawk (Dare County) (Fig. 1). Most

sightings have been of single individuals; the largest groups sighted

for several days before disappearing were three at Minnesott Beach

Yacht Basin, a tributary to the Neuse River (Pamlico River) in Au-

gust 1980, and three in the Trent River at the Sheraton Yacht Harbor,

New Bern (Craven County) in September 1994.

Although six records are of manatees frequenting Pelletier Creek,

the area seems an unlikely manatee habitat as the short 0.8 km creek

is plied by many boats, pollution is heavy from numerous boat works

and marinas, and considerable runoff occurs from the densely built

houses and condominiums lining the shoreline. Yet manatees have

spent several days in the creek during each visit. In 1993 to 1994 two

(once) to six separate manatees frequented the same general area of

the coffee-colored Trent River, tributary to the Neuse River, 2 km

southwest of New Bern (Craven County) and fed on dense aquatic

vegetation. Three specimens frequented the river 0.2 km west of New

Manatees in North Carolina 59

Bern, at Lawson Park or the Sheraton Hotel Yacht Basin in 1994. The

1993 specimen was followed downstream in the Neuse along its

western shore for 13 km before vanishing. The July 1994 specimen

did not linger in the Trent River prior to its next (presumed) sight-

ing in the lower Neuse River opposite Oriental in late July. August-

September 1994 specimens lingered and fed on the lush vegetation of

the Trent River even as late as 24 September 1994. Bottom and surface

salinities 17 September 1994 were 14 and 7 ppt respectively.

INLAND PENETRATIONS

Manatees are known to penetrate inland freshwater such as the

St. John's River of Florida for 224 km to Lake Monroe (Volusia

County; D. Odell, Sea World Inc., personal communication). Farthest

inland river penetrations by manatees in North Carolina have been:

Cape Fear River (one) for 94.4 km-6.4 km above Route 210 and north

of Wilmington (Pender County) 11 July 1993, and one for 92 km in

the Neuse River to the Fort Barnwell Bridge (Craven County) in

October 1980, 33 km northeast of New Bern (Fig. 1). Two other

manatees occurred in the Tar River system of the upper Pamlico

River, a tributary of Pamlico Sound: one for 88 km to Greenville (Pitt

County) in November 1989 and the other for 58 km to just upstream

of Washington (Beaufort County) (Fig. 1) in September 1985. The

most peculiar movements have been of three manatees that traversed

south from the lower Chesapeake Bay in late summer of 1993 via the

canal and Intracoastal Waterway into Currituck Sound (Fig. 1). While

most manatee sightings have been at localities which could have been

reached via the Intracoastal Waterway and sounds, the four inlet and

six ocean occurrences also suggest travel north from Florida to North

Carolina may have been via those avenues. In any event, observations

seem to indicate many more young manatees are expanding their range

into North Carolina, perhaps as a result of an increased public aware-

ness of Florida's manatees rather than a real population increase.

ACKNOWLEDGMENTS— Thanks are extended to the many

interested people of North Carolina who called or commented on

manatee occurrences in North Carolina. Others, elsewhere, who were

also helpful in supplying information were: C. Beck, National Bio-

logical Service, Sirenia Project, Gainesville, FL (occurrences, sizes,

and Rathbun et al. 1981 reference); D. Odell, Sea World Inc., Or-

lando, FL, (St. John's River occurrence, file data, and review of manu-

script; R. Dove, Neuse River Keeper (1994 Trent River observations);

J. Wolfenberger, (Minnesott Beach Yacht Basin observations); New

60 Frank J. Schwartz

Hanover Public Library (Cape Fear River stories); J. Oakley, Carolina

Biological Lab, Burlington (early Brimley references); M. Nabinetts,

Washington Daily News (Pamlico River stories), N. Winfrey, Orien-

tal News (comments on lower Neuse River observations); M. Clark

provided the three North Carolina State Museum of Natural Sciences

records and checked the Museum overall occurrence records. D. Lee

of the North Carolina State Museum of Natural Sciences reviewed the

paper, as did C. Potter, National Marine Fisheries Service, Division

of Mammals (North Carolina data). D. Webster, University of North

Carolina, Wilmington, noted the 29 July 1992 record. L. White typed

the text; R. Barnes produced Figure 1.

LITERATURE CITED

Brimley, C. S. 1905 (1908). A descriptive catalogue of the mammals

of North Carolina, exclusive of the Cetacea, Journal of the Elisha

Mitchell Scientific Society 21:1-32.

Brimley, C. S. 1946. The mammals of North Carolina, installment

No. 17. Carolina Tips 9(l):2-3.

Brimley, H. H. 1931. Mammals of North Carolina. Journal of Mammalogy

12:320-321.

Caldwell, D. K., and F. B. Golley. 1965. Marine mammals from the

coast of Georgia to Cape Hatteras. Journal of the Elisha Mitchell

Scientific Society 81:24-32.

Campbell, H. W. 1977. Mammalia: Sirenia; Trichechidae {Tricheches

manatus Linnaeus). Pages 396-397 in Endangered and threatened

plants and animals of North Carolina (J. E. Cooper, S. S. Robinson,

and J. B. Funderberg, editors). North Carolina State Museum of

Natural History, Raleigh.

Jefferson, T. A., S. Leatherwood, and M. A. Webber. 1993. Marine

mammals of the world. FAO Species Identification Guide, Rome,

Italy.

Lee, D. S. 1976. Manatees: Mammals in distress. Wildlife in North

Carolina 40(7):8-9, 27.

Mitchell, K. B. 1993. My god, that's not a beaver. Wilmington Star

News, 20 July: la, 4a.

Rathbun, G. B., R. K. Bonde, and D. Clay. 1981. The status of the

West Indian manatee on the Atlantic Coast north of Florida. Pages

152-165 in Proceedings of the Symposium on nongame and en-

dangered wildlife (R. R. Odom and J. W. Guthrie, editors). Georgia

Department of Natural Resources, Game Fish Division Technical

Bulletin WL5, Atlanta.

Smith, E. R., J. B. Funderberg, and T. L. Quay. 1980. A checklist

of North Carolina mammals. North Carolina Wildlife Resource

Commission, Raleigh.

Received 13 October 1994

Accepted 10 February 1995

Recovery of the Cave Crayfish (Decapoda: Cambridae)

Population in Peacock Springs, Florida?

W. J. Streever

Department of Environmental Engineering Sciences,

University of Florida, Gainesville, Florida 32611

ABSTRACT— In 1991, a cave fauna kill was observed within

the fully-flooded cave system at Peacock Springs, Suwannee

County, Florida. Organisms affected by the kill included the

pallid cave crayfish (Procambarus pallidus), the yellow bull-

head (Ameiurus natalis), the American eel (Anguilla rostrata),

and the Asiatic clam (Corbicula fluminea). Following the kill,

no crayfish were present along transects, but recently collected

census data are not significantly different from data collected

before the 1991 kill (Paired Mest, P > 0.1). However, cray-

fish numbers have not returned to their pre- 1991 levels in the

cave passage where the highest crayfish density occurred

before the kill. Also, the scarcity of small (<1.5 cm total

length) crayfish suggests that the return to pre-1991 levels may

reflect dispersal of animals from inaccessible portions of the

cave and not replacement of crayfish through reproduction.

The troglobitic pallid cave crayfish, Procambarus pallidus,

inhabits flooded caves in the Suwannee River drainage in north

Florida (Franz and Lee 1982). These caves also are inhabited by other

troglobitic and troglophillic species, including several species of

catfish (Hale and Streever 1994), ostracods (Walton and Hobbs 1959),

the American eel Anguilla rostrata (Franz et al. 1994), the Asiatic

clam Corbicula fluminea (Streever 1992a), tubificidae worms (Streever

19926), and the colonial cnidarian Cordylophora lacustris (Streever

1992c). Franz et al. (1994) provide an extensive review of Florida's

cave fauna, emphasizing distributional records for all species report-

ed in association with caves. Little is known about the ecology or

population dynamics for most of these species.

The Peacock Springs cave system consists of over 3,300 m of

fully-flooded passage, with depths down to about 60 m. Eight openings

provide access to the cave system. A surface channel connects open-

ings commonly called Peacock I, II, and III to the Suwannee River.

Water from the Floridan aquifer normally flows out of the Peacock I

opening and drains into the Peacock III opening and the Suwannee

Brimleyana 22:61-65, June 1995 61

62 W. J. Streever

River. However, during periods of high river levels, water from the

Suwannee River overcomes the head of pressure generated by aquifer

water. Flow in the surface channel reverses, and Suwannee River

water enters the cave system. Following a flow reversal in February

1991, virtually all of the animals in accessible portions of the Peacock

Springs cave system were killed. Animals affected by the kill include

crayfish, catfish, and Asiatic clams (Streever 19926). The cause of

the kill is unknown.

This paper addresses two issues. First, I present 1994 P. pallidus

census data and subjective observations of other taxa, and second

I discuss a possible mechanism for the increase in crayfish population

density following the 1991 kill.

METHODS

In 1990, divers established eight belt transects (100 x 4 m) in

association with permanent guidelines in the Peacock Springs cave

system. Transects were numbered 1-8. Both before and after the fauna

kill in February 1991, crayfish were visually censused along transects

by divers carrying 50-watt lights. Because cave crayfish roam in the

open on the cave floor, visual censusing provides a simple method of

estimating population density. After the kill, two additional belt tran-

sects, called 2-a (240 m long) and 8-a (350 m long), were established

in a tunnel north of the Peacock I opening and north of the Peacock

III opening, respectively. Transect 2-a incorporated transect 2, and

transect 8-a incorporated transects 7 and 8. All crayfish that were

visible from the cave's permanent guidelines were included in censuses

along transects 2-a and 8-a, making their width dependent on water

clarity and size of the cave passage. In general, the width of transects

2-a and 8-a varied between 4 and 10 m. The large size of the new

transects was intended to allow censusing of the smaller crayfish

densities occurring after the 1991 kill. A map of the cave showing

transect locations was provided by Streever (19926).

The 1994 transect censuses were made between 9 January 1994

and 25 November 1994. During the 1994 censuses, crayfish were

recorded as belonging to a size class of less than or greater than 1.5

cm in overall length, estimated from the anterior edge of the rostrum

to the posterior edge of the telson. All censuses along each of the

original transects were averaged to yield values for each transect

before the kill and in 1994. The null hypothesis of no difference

between censuses taken before the 1991 kill and those taken in 1994

was tested by a paired Mest, with averages of before and after censuses

along individual transects comprising pairs (Zar 1984).

Crayfish Population 63

RESULTS AND DISCUSSION

Differences in censuses (Table 1) before the 1991 kill and in

1994 are not statistically significant (P > 0.1). The increase in crayfish

population density immediately after the kill and in 1994 suggests

that the population is recovering. However, 1994 censuses in the Peacock

III tunnel indicate that populations have not recovered to pre-kill

levels along transects 7 and 8. Transect 8a, which encompassed transects

7 and 8 as well as an additional 100 m of cave passage, contained

only a fraction of the number of crayfish that were present along

transects 7 and 8 before the kill. This indicates that the large crayfish

population found in the Peacock III tunnel before the kill is not

recovering, nor is the overall population in the cave system.

Of the 138 crayfish censused after the 1991 kill, only one

animal was smaller than 1.5 cm. Although animals smaller than 1.5

cm in total length may be missed during censusing, the scarcity of

Table 1. Census data for Procambarus pallidus in the Peacock Springs,

Florida, cave system before the 1991 kill, immediately after the kill,

and in 1994.

These transects were established after the 1991 kill to cover a larger area than

original transects.

64 W. J. Streever

small animals suggests that recovery of crayfish population density is

not dependent on replacement through reproduction. Cave popula-

tions frequently display size distributions with many large individuals

and few small individuals, suggesting infrequent reproduction (Poulson

1963, Culver 1982). The fecundity of P. pallidus is unknown, but an

extensive study of the cave-adapted crayfish Orconectes australis

australis in Shelta Cave, Alabama, suggests that reproduction in cave

crayfishes may be infrequent and that clutch sizes may be small

(Cooper 1975). If a similar pattern occurs in P. pallidus, crayfish

population recovery that depends on reproduction could be slow. Be-

cause crayfish population densities in the Peacock Springs cave sys-

tem do not appear to be recovering through reproduction, the increase

in numbers along transects 1 through 6 may not represent a true

recovery of the crayfish population. Instead, crayfish density may be

the result of colonization by individuals that survived the kill and

are now moving into the portion of the cave where transects were

located.

Catfish were not counted as part of this study. However, they

were common along transects before the 1991 kill, they were absent

from transects immediately after the kill, and they were common

during the 1994 census dives. An Asiatic clam population near the

mouth of Peacock III had an estimated density of 161 individuals/m 2

before the 1991 kill (Streever 1992a), but no live Asiatic clams were

found immediately after the kill or in 1994. Empty clam shells, pre-

sumably remaining from before the kill, were abundant.

The causes of the kill and the factors affecting recovery are

unknown. However, intrusion of river water into caves typically full

of groundwater appears to be linked to cave kills in the Peacock

system. On 19 November 1994, two dead crayfish and a dead catfish

were found along transect 6, immediately after a period of flooding

during which Suwannee River water entered the cave. However, seven

live crayfish were found along the transect on the same day, so this

event did not have the intensity of the 1991 kill. Despite this apparent

association, the link between river water and crayfish mortality is not

clear. P. pallidus has been observed in two siphon-spring cave

systems flooded by Sante Fe River water throughout the year

(personal observation), so intolerance of low temperatures and other

environmental factors associated with river water may not be the

cause of mortality. Furthermore, kills have been observed in other

caves without flooding by river water (Franz et al. 1994). Establish-

ment of the cause of cave crayfish kills requires further investigation.

Crayfish Population 65

ACKNOWLEDGMENTS—! thank cave divers Jason Hale, Wendy

Short, Steve Brooker, and Rob Korn for their assistance with field

work. The State of Florida Department of Parks and Recreation allowed

access to the Peacock Springs cave system. This work was funded in

part by a National Science Foundation Graduate Research Fellowship.

LITERATURE CITED

Cooper, J. E. 1975. Ecological and behavioral studies in Shelta

Cave, Alabama, with emphasis on decapod crustaceans. Ph.D. Thesis.

University of Kentucky, Lexington.

Culver, D. C. 1982. Cave life, evolution and ecology. Harvard Uni-

versity Press, Cambridge, Massachusetts.

Franz, R., J. Bauer, and T. Morris. 1994. Review of biologically

significant caves and their faunas in Florida and south Georgia.

Brimleyana 20:1-109.

Franz, R., and D. S. Lee. 1982. Distribution and evolution of Florida's

troglobitic crayfishes. Bulletin of the Florida State Museum Bio-

logical Sciences 28:53-78.

Hale, J. A., and W. J. Streever. 1994. Cave fauna distribution

within fully-flooded cave systems in Florida. Journal of Freshwa-

ter Ecology 9:171-174.

Poulson, T. L. 1963. Cave adaptation in amblyopsid fishes. Ameri-

can Midland Naturalist. 70:257-290.

Streever, W. J. 1992a. First record of Corbicula clams within flooded

cave systems. Florida Scientist 55:35-37.

Streever, W. J. 1992/?. Report of a cave fauna kill at Peacock

Springs cave system, Suwannee County, Florida. Florida Scientist

55:125-128.

Streever, W. J. 1992c. First record of the colonial cnidarian Cordylophora

lacustris within a flooded cave system. NSS Bulletin 54:77-78.

Walton, M., and H. H. Hobbs, Jr. 1959. Two new eyeless Ostracods

of the genus Entocythere from Florida. Quarterly Journal of the

Florida Academy of Sciences 22:114-120.

Zar, J. H. 1974. Biostatistical analysis. Prentice-Hall, Englewood

Cliffs, New Jersey.

Received 1 January 1995

Accepted 14 February 1995

Premolar Cementum and Noncementum Lengths

As Potential Indicators of Age for Beavers,

Castor canadensis (Rodentia: Castoridae)

Allan E. Houston

Ames Plantation, Agricultural Experiment Station, The University

of Tennessee, P. O. Box 389, Grand Junction, Tennessee 38039-0389

AND

Michael R. Pelton

Department Forestry, Wildlife, and Fisheries, The University

of Tennessee, Knoxville, Tennessee 37901-1071

ABSTRACT — Jaws from 28 beavers (Castor canadensis), >4 years

old as determined from cementum annuli on premolars and

molars, were collected in west Tennessee. An exponential model

predicting age was developed based on the independent vari-

ables cementum length and noncementum length as measured

along the estimated, maximum longitudinal centerline of cross-

sectioned premolars: AGE = 0 + 6.1784 * e (:o.io3i*noncemen) +

2 6513 * e (0.1U9*CEMENTUAf)

Several methods of aging beavers have been attempted with varying

degrees of accuracy, including pelt size (Buckley and Libby 1955),

tail dimensions, total body mass, skull measurements (Patric and Webb

1960), and baculum size and mass (Friley 1949). Probably the most

accurate method was developed by van Nostrand and Stephenson (1964)

using tooth eruption and closure of basal openings of premolars and

molars for beavers up to 3 years old, and cementum layering in premolars

and molars for specimens >4 years. Larson and van Nostrand (1968)

further refined this technique to include criteria dealing with cementum

deposition around basal openings. They noted that age estimations

may be complicated by multi-annual cementum layering, but that the

ratio of cementum to noncementum on molars and premolars might

be used to estimate age in older specimens.

Our objective was to use premolar cementum length (aggregate

of all annual cementum depositions) and noncementum length (remains

of the original tooth) as independent variables to evaluate a model for

estimating age of beavers >4 years.

Brimleyana 22:67-72, June 1995 67

68 Allan E. Houston and Michael R. Pelton

METHODS

This study was conducted in the upper headwater basin of the

North Fork of the Wolf River in west Tennessee, largely on the Ames

Plantation. From November 1984 through May 1985, beaver jaws

from 169 beavers were collected from legal trapping efforts on

approximately 1,619 ha of the watershed.

Molars and premolars from lower mandibles were extracted and

cleaned. If tooth basal openings indicated a specimen to be >4 years

old, age was determined by grinding the lingual surface of premolars

or molars with a 120-grit stone to expose longitudinal cross-sections.

Ground surfaces were polished with a 400-grit emery cloth and cementum

layers were carefully counted using hand-held magnification (Larson

and van Nostrand 1968). Twenty-eight beavers, >4 years old as aged

in this manner (Table 1), provided the samples (i.e., dependent variables)

for our study.

Premolar cementum length and noncementum length were measured

to the nearest millimeter along the estimated maximum longitudinal

cross-sectional centerline of one premolar per specimen (Fig. 1). An

exponential model predicting age in years was developed by combining

a growth curve for cementum length and a decay curve for noncementum,

using nonlinear regression (PROC NLIN) and the multivariate secant

method (DUD) to set initial parameters (SAS Institute, Inc. 1985).

Intercept was specified at zero to prevent predicted ages from

dipping below zero. An "r2 like" statistic was calculated by taking

[1 - (residual SS/corrected total SS)].

RESULTS AND DISCUSSION

The following model was developed using cementum length and

noncementum length of premolars as independent variables:

AGE = 0 + 6 1784 * e (-o.io37*noncemen) + 2.6513 * e (°ul9*CEMENTUM)t

The "r2 like" statistic was 0.93. The predicted curves are combined

to produce a response surface (Fig. 2).

Our data were obtained from specimens on the headwaters of a

single watershed. Although beavers can travel considerable distances,

we assumed our sample represented only a small region. The study

area may not fully represent variation existing range-wide or within

adjacent watersheds. It is possible that genetic differences and dietary

regimes will yield different tooth size, wear, and cementum accretion

Aging Beaver Teeth 69

Table 1. Age, as estimated by cementum annuli, cementum and noncementum

length, measured to the nearest millimeter along the maximum longitu-

dinal cross-section of one premolar per specimen, and predicted ages

and residuals for beavers captured in west Tennessee, November 1984

through May 1985.

AGE = 6.174*e<-°-1037*"°"C£M£") + 2.6513V0

1U9*CEMENTUM)

70

Allan E. Houston and Michael R. Pelton

Fig. 1. Schematic representation of cementum length and noncementum

length as measured along longitudinal cross-sections of premolars for

beaver.

Aging Beaver Teeth

71

PREDICTEDAGE

(YEARS)

V 12.00

CEMENTUM LENGTH

67 (MM)

18.67

NONCEMENTUM LENGTH

(MM)

Fig. 2. Response surface of an exponential model predicting age of

west Tennessee beaver using cementum length and noncementum length

measured along the estimated maximum longitudinal cross-sections of

premolars as independent variables.

patterns. Comparative study is needed to document potential variation

of these criteria. Also, the method should be validated using known-

age specimens. However, these results suggest that this technique

could be developed as a reliable method to age beavers.

ACKNOWLEDGMENTS— -The project was funded by the Hobart

Ames Foundation and The University of Tennessee Agricultural Experiment

Station. J. Schneider gave invaluable assistance with statistical analyses.

R. Henry, J. Morrow, and T. Sain provided the majority of data collections.

R. Maxey contributed the drawing in Figure 1.

LITERATURE CITED

Buckley, J. J., and W. L. Libby. 1955. Growth rates and age determi-

nation in Alaska beaver. Transactions of the North American

Wildlife and Natural Resources Conference 20:495-505.

Friley, C. E., Jr. 1949. Use of the baculum in age determination of

Michigan beaver. Journal of Mammalogy 30(3):261-267.

Larson, J. S., and F. C. van Nostrand. 1968. An evaluation of beaver

aging techniques. The Journal of Wildlife Management 32(1):99-103.

72 Allan E. Houston and Michael R. Pelton

Patric, E. F., and W. L. Webb. 1960. An evaluation of three age

determination criteria in live beavers. The Journal of Wildlife

Management 24:37-44.

SAS Institute, Inc. 1985. SAS User's Guide: Statistics. SAS Institute,

Inc., Box 8000, Cary, North Carolina,

van Nostrand, F. C, and A. B. Stephenson. 1964. Age determination

for beavers by tooth development. The Journal of Wildlife

Management 28(3):430-434.

Received 11 January 1994

Accepted 8 March 1995

Record Clutch Size for Chelydra serpentina

(Testudines: Chelydridae) in Virginia

Joseph C. Mitchell

Department of Biology

University of Richmond,

Richmond, Virginia 23173

AND

Michael C. Odom

U.S. Fish and Wildlife Service

11110 Kimages Road

Charles City, Virginia 23030

ABSTRACT — A record clutch size of 67 is reported for a

common snapping turtle {Chelydra serpentina serpentina) from

Charles City County, Virginia, 12 larger than previously

reported for this chelonian. The new average for this species

in the Commonwealth is 29.9 ± 16.6 ( x ± SD).

The snapping turtle (Chelydra serpentina) occurs from

Florida north to southern Canada and Nova Scotia in eastern North

America and from the Mexican border north to extreme southern

Saskatchewan in western North America (Conant and Collins 1991,

Iverson 1992, Russell and Bauer 1993). Clutch size varies latitudinally,

with the largest sizes reported for northern populations in Canada,

South Dakota, and Nebraska (Zug 1993; J. B. Iverson, Earlham College,

personal communication). Ash (1951) reported an average clutch

size of 28.6 and a range of 13-48 for 85 specimens presumably caught

in the Virginia area. However, these data were presented only in

an abstract. A complete paper was never published, and no docu-

mentation is available for us to determine the source of his

specimens. Information provided by the late John T. Wood

(formerly a retired medical doctor in Victoria, British Columbia

and Virginia in the 1950s, personal communication) indicated that

Ash's specimens could have been from multiple locations in and

out of Virginia. Mitchell (1994) reported an average clutch size of

27.0 ± 13.2 (range = 7-55) for 13 females collected from various

locations in Virginia.

On 3 June 1994, a gravid female (271-mm carapace length,

201-mm plastron length) was inadvertently killed by a vehicle

at Harrison Lake National Fish Hatchery, Charles City County,

Virginia. Subsequent dissection revealed 67 shelled eggs. One egg

Brimleyana 22:73-75, June 1995 73

74 Joseph C. Mitchell and Michael C. Odom

was opened on the date of collection, and the rest were placed in a

closed plastic container with a 3:1 mixture of top soil and vermiculite.

The soil was kept moist during the incubation period so that the

relative humidity was maintained at nearly 100%. Incubation temperatures

varied from 25 to 30° C. During the course of incubation, 12 of the

66 eggs died and were discarded; 54 eggs survived. Hatching

commenced on 20 August (78 days incubation), and the last egg

pipped on 24 August (82 days). Hatchlings showed considerable

variation in development, with larger individuals bearing a small

yolk sac and smaller individuals bearing a large, cumbersome yolk

sac. Of the 54 hatchlings, four failed to survive through yolk sac

absorption. The remaining turtles began exogenous feeding as their

yolk sac diminished, taking a diet of chopped nightcrawlers and

miscellaneous live aquatic macroinvertebrates collected from

ponds. The surviving juvenile turtles were released the following

spring.

A documented clutch size of 67 eggs is 12 larger than the

largest reported for Virginia (Mitchell 1994). Incorporation of the

large clutch size reported here yields a new state average of 29.9 ±

16.6 (range = 7-67). The maximum known clutch size for Virginia

populations lies between those known for northern populations

(83: Quebec (Bleakney 1957); 73: New York and Wisconsin (Yntema

1970); 109: Nebraska (Packard et al. 1990)) and southern populations

(43: North Carolina (Brown 1992); 21: Florida (Punzo 1975)).

Reports of average clutch sizes for snapping turtles in different

geographic areas based on small sample sizes should be used with

caution. Such averages will almost always change with additional

data.

ACKNOWLEDGMENTS— Viz thank Kurt A. Buhlmann and John

B. Iverson for their comments on the manuscript, and Thomas F.

Burnett for assistance in the collection of the adult turtle and

maintenance of the eggs and hatchlings.

LITERATURE CITED

Ash, R. P. 1951. A preliminary report on the size, egg number, incu-

bation period, and hatching in the common snapping turtle, Chelydra

serpentina. Virginia Journal of Science 2:312 (abstract).

Bleakney, S. 1957. A snapping turtle, Chelydra serpentina serpentina,

containing eighty-three eggs. Copeia 1957:143.

Chelydra serpentina clutch 75

Brown, E. E. 1992. Notes on amphibians and reptiles of the western

Piedmont of North Carolina. Journal of the Elisha Mitchell Sci-

entific Society 108:38-54.

Conant, R., and J. T. Collins. 1991. A field guide to the reptiles

and amphibians of eastern and central North America. Houghton

Mifflin Company, Boston, Massachusetts.

Iverson, J. B. 1992. A revised checklist with distribution maps of

the turtles of the world. Privately Printed, Richmond, Indiana.

Mitchell, J. C. 1994. The reptiles of Virginia. Smithsonian Institution

Press, Washington, D.C.

Packard, G. C, M. J. Packard, and K. Miller. 1990. Life history:

Chelydra serpentina, fecundity. Herpetological Review 21:92.

Punzo, F. 1975. Studies on the feeding behavior, diet, nesting

habits and temperature relationships of Chelydra serpentina osceola

(Chelonia: Chelydridae). Journal of Herpetology 9:207-210.

Russell, A. P, and A. M Bauer. 1993. The amphibians and reptiles of

Alberta. University of Calgary Press, Calgary, Alberta, Canada.

Yntema, C. L. 1970. Observations on females and eggs of the com-

mon snapping turtle Chelydra serpentina. American Midland Naturalist

84:68-76.

Zug, G. R. 1993. Herpetology, an introductory biology of amphibians and

reptiles. Academic Press, San Diego, California.

Received 12 January 1995

Accepted 9 March 1995

New Distributional Records for the Star-nosed Mole,

Condylura cristata (Insectivora: Talpidae), in

North Carolina, with Comments on its Occurrence

in the Piedmont Region

Jeffrey C. Beane

North Carolina State Museum of Natural Sciences

P.O. Box 29555

Raleigh, North Carolina 27626-0555

ABSTRACT — The distribution of the star-nosed mole (Condylura

cristata) in North Carolina is updated, including specimen-supported

records for seven new counties and sight records for several

additional counties. The species is locally common in the Mountains,

spottily distributed throughout much of the Coastal Plain, and

apparently rare to absent in much of the Piedmont. A specimen

from just off the Blue Ridge escarpment in western Surry

County represents the first Piedmont record supported by a

specimen, although there are additional sight records and unverified

reports from that region. This mole is sympatric with the state's

other two mole species in many areas, but usually occurs in

wetter habitats than either.

INTRODUCTION

The star-nosed mole (Condylura cristata) ranges from eastern

Manitoba and Minnesota eastward to Labrador and Nova Scotia, and

southeastward along the Atlantic coast to southeastern Georgia (Paradiso

1959, Burt and Grossenheider 1976, Petersen and Yates 1980, Hall

1981, Yates and Pedersen 1982). In North Carolina, its distribution

has frequently been reported as consisting of two distinct popula-

tions— one in the Mountains, where the species is locally common,

and one in the Coastal Plain, where it is uncommon to rare — with a

conspicuous hiatus in the Piedmont (Lee et al. 1982, Webster et al.

1985, Lee 1987, Webster 1987). The scarcity of records from the

Coastal Plain and complete absence of Piedmont records resulted in

the species being recommended for listing as status Undetermined by

the Nongame Advisory Committee to the North Carolina Wildlife

Resources Commission (Webster 1987), and in 1990 the species was

granted protection as a species of Special Concern under the North

Carolina Endangered Species Act (G.S. 113-331 to 113-337).

I here report on the current known distribution of Condylura

cristata in North Carolina by providing a list of existing records from

the state, and a dot distribution map (Fig. 1) updated from maps

provided by Lee et al. (1982) and Webster (1987).

Brimleyana 22:77-86, June 1995 77

78

Jeffrey C. Beane

Fig. 1. Known distribution of Condylura cristata in North Carolina.

Solid dots indicate localities documented by specimens in curated collections.

Open circles indicate literature records or sight records not supported

by specimens. A single dot or circle may represent two or more records

in close proximity. A "C" indicates a literature record, not supported

by a specimen, for which only the county and no further information

was given. Question marks accompany records for which no specimen

was observed, or which are unverifiable or otherwise questionable.

RECORDS OF OCCURRENCE

The following North Carolina localities are supported by voucher

specimens, many of them in the North Carolina State Museum of

Natural Sciences (NCSM), and are represented by solid dots on the

distribution map:

Alleghany Co.: New River, "Sparta" (The New River is actually, at

its nearest point, 4 mi. NW of Sparta), 30 May 1968, H. M. Tyus

(NCSM 1762). Ashe Co.: "3A mi. from jet. NC 16 and Friendship

Church Rd." (= ca. 2 mi. ENE Jefferson), 4 Oct. 1991, E. Marsh

(Appalachian State University 16731). Avery Co.: 2.6 mi. NNE Heaton,

Pisgah National Forest, 27 May 1975, R. Jordan and J. Pentecost

(Auburn University Museum 1-281); 3A mi. SE Linville, 24 July 1984,

D. S. Lee and J. P. Kumhyr (NCSM 4875). Brunswick Co.: ca. 6 mi.

NNW Supply, Green Swamp Ecological Preserve off NC 211, "Moon

Island." 16 May 1993, T. M. Padgett (TMP 511; to be deposited at

NCSM). Buncombe Co.: W of Asheville (University of Michigan Mu-

seum of Zoology); Biltmore (Harvard Museum of Comparative Zool-

ogy 6635); ca. 2.6 mi. WNW Swannanoa, Warren Wilson College

campus, Aug. 1977, D. S. Lee and B. Lee (NCSM 7837); Weaverville,

6 Feb. 1896, Mrs. J. S. Cairns (Harvard Museum of Comparative

Zoology 5399). Dare Co.: ca. 7 mi. SW Manns Harbor, Alligator

Star-nosed Mole 79

River National Wildlife Refuge, Milltail Rd., 20 March 1991, P. R.

Trail, M. L. Dunn, et al. (NCSM 6710). Haywood Co.: 5 mi. SW

Waynesville on U.S. 23, 30 Aug. 1951, F. S. Barkalow, Jr. (North

Carolina State University 274). Henderson Co.: Hoopers Creek near

Fletcher, 8 Oct. 1974, B. A. Sanders (NCSM 4937); 1 mi. S Gerton,

Little Bearwallow Mtn., 8 July 1994, P. B. Spivey (to be deposited at

NCSM); 1 mi. W. NW Mills River (town), near jet. SR 1336 and SR

1338, 25 May 1993, K. A. Buhlmann, D. W. Herman, and J. L. Warner

(NCSM 7868). Hoke Co.: 9.2 mi. NE Ashley Heights, Ft. Bragg Mili-

tary Reservation, 29 May 1992, LCTA survey crew (NCSM 7213).

Jackson Co.: 0.7 mi. NE Balsam, Blue Ridge Parkway at Balsam

Gap, 22 May 1991, A. C. Boynton (NCSM 7528). Macon Co.: High-

lands, 1 June 1946 (University of Georgia Museum of Natural History

192); Highlands, ca. 1 June 1946, R. Bridges (NCSM 6575); ca. 0.5

mi. E center Highlands, Highlands Biological Station, 23 Oct. 1982,

S. Morrison (University of Georgia Museum of Natural History 7595);

1 mi. E.NE Highlands, SR 1603, 0.3 rdmi. E jet. SR 1604, 20 Sept.

1984, A. L. Braswell and J. E. Cooper (NCSM 7462). New Hanover

Co.: near Carolina Beach, 16 Nov. 1956, G. Tregembo (NCSM 3243).

Mitchell Co.: Roan Mtn. (Wake Forest University); Roan Mtn. near

Magnetic City, 15 March 1894 (U. S. National Museum 66300); Roan

Mtn. near Magnetic City, 15 Aug. 1895 (U. S. National museum

91519). Polk Co.: Green River Valley near Saluda, 28 May 1974,

G. G. Shaw and M. Bradley (NCSM 1237). Rutherford Co.: Chimney

Rock, Southside Drive, 19 Dec. 1989, M. Jaeger-Gale (NCSM 6336).

Richmond Co.: 3.7 mi. NE Hoffman, U.S. 1, 0.1 rdmi. S Lumber

River, 18 May 1980, A. L. Braswell (NCSM 3283). Sampson Co.: 5.5

km W Faison (Duplin Co.) off SR 1734, 9 June 1994, W. D. Webster

(University of North Carolina at Wilmington 3370). Scotland Co.:

"Laurinburg, Mr. Newton's farm" [the farm referred to was located

ca. 8 mi. N Laurinburg, or 4.5 mi. W Wagram (NCSM files, W. M.

Palmer, pers. comm.)], 12 April 1975, R. B. Julian (NCSM 3047).

Surry Co.: 0.8 mi. NNW Low Gap near jet. NC 89 and SR 1433,

A. B. Somers et al., 7 June 1994 (NCSM 7744). Swain Co.: Deep

Creek; Kephart Prong Hatchery; Smokemont; Charlies Bunion Mtn.

at Sevier Co., TN line; and Appalachian Trail between Newfound

Gap and Indian Gap at Sevier Co., TN line; all in Great Smoky

Mountains National Park (Great Smoky Mountains National Park

collection, Linzey and Linzey 1968); Great Smoky Mountains

National Park, Appalachian Trail, lA mi. S Boulevard Trail (= ca. 6.8

mi. NW Smokemont), 20 Aug. 1961, J. B. Westbrook (University of

Georgia Museum of Natural History 3112). Watauga Co.: Blowing

80 Jeffrey C. Beane

Rock (Wake Forest University); Boone (Appalachian State University

7606); Boone, Hidden Valley Circle, 14 March 1981, D. Notrichia

(Appalachian State University 7591); Boone, 1987, R. W. Van Devender

(Appalachian State University 14633); ca. 1 mi. WSW Boone, jet.

NC 105 and Poplar Grove Rd., 16 Sept. 1978 (Appalachian State

University 5125 ); ca. 3 mi. WSW Boone, Laurel Cr. along NC 105,

13 Sept. 1985, F. Authenreith (Appalachian State University 12086);

Blue Ridge TWP, Jakes Mtn. Rd. near jet. SR 1511, Dec. 1986, M. P.

Rowe (Appalachian State University 12786); Blue Ridge TWP, SR

1511 along Sand Spit Branch and Blue Ridge Parkway, 27 Sept. 1987,

R. W. Van Devender (Appalachian State University 14205); Sugar

Grove, jet. U.S. 321 and Edmiston Rd., 24 Nov. 1978, H. Rogers

(Appalachian State University 7601); Sugar Grove, Edmiston Rd., 5

Sept. 1985, R. N. Henson (Appalachian State University 12485); Watauga

Co., no further data (Appalachian State University 16751). Yancey

Co.\ Burnsville, 3 Feb. 1976, L. P. Hartis (NCSM 2409).

In addition to localities supported by voucher specimens, the

following literature and sight records are considered valid, and most

are included as open circles on the distribution map:

"Mountains near the border of South Carolina" (Audubon and Bachman

1851). Avery Co.: Elk Park (Lee 1987, NCSM files). Bladen Co.: ca.

2 mi. N.NE White Lake (town) off SR 1517 near jet. U.S. 701, 30

April 1992, T. M. Padgett and R. Rageot (Thomas M. Padgett, pers.

comm.). Buncombe Co.: 2-3 mi. N Black Mountain at Montreat, ca.

1915-1920, E. E. Brown (Elmer E. Brown, pers. comm.); Swannanoa,

ca. 20 April 1931, C. S. Clapp (Lee 1987, NCSM files). Cherokee

Co.: Topton, 13 May 1934, E. B. King (Brimley 1944-1946). Clay

Co.: Hayesville, 3 Aug. 1947, R. G. Vick (Paradiso 1959, NCSM

files; specimen formerly at NCSM but apparently lost). Macon Co.:

Highlands vicinity, specimens "not infrequently" taken by H. C. Harbison

prior to May 1908 (Brimley 1944-1946); Highlands vicinity, three

specimens reported by Johnston (1967); one of these, formerly in the

Highlands Museum, is now NCSM 6575; the same specimen was also

reported by Odum (1949). McDowell Co.: just N Ashford along U.S.

221, ca. 1800' elev., two specimens taken from the stomach of a

black rat snake (Elaphe obsoleta), 23 May 1960, E. E. Brown. These

specimens were reported by Brown (1979), but their specific locality

was not included in that paper (Elmer E. Brown, pers. comm.). Mecklenburg

Co.: Davidson (east side near golf course branch), ca. 1950-1955, E.

E. Brown (Elmer E. Brown, pers. comm.). New Hanover Co.: near

Carolina Beach, 7 June 1959, G. Tregembo (Lee 1987, NCSM files).

Pitt Co.: no further data (Lee et al. 1982, Clark et al. 1985, Lee 1987,

Star-nosed Mole 81

Webster 1987). The original source of this record seems uncertain,

and it is accompanied by a question mark on the distribution map in

Fig. 1. Robeson Co.: near Lumberton, early Dec. 1943, A. M. Ivey

(Brimley 1944-1946). Sampson Co.: near Garland, 13 May 1918, J. F.

Johnson (Brimley 1923, Brimley 1944-1946, Paradiso 1959). Washington

Co.: Wenona, early Nov. 1992, E. R. Rainey (Brimley 1923, Brimley

1944-1946, Paradiso 1959). Watauga Co.: Boone, four specimens, 19

May 1918, (Brimley 1944-1946). Yancey Co.: ca. 7 mi. W. Burnsville

along Lickskillet Branch, ca. 1981. J. McFee (Allen C. Boynton,

pers. comm.).

Lee (1987) and Webster (1987) reported C. cristata from Moyock

in Currituck County, based on a specimen from the digestive tract of

a mink obtained from a fur dealer in that town and reported by Wilson

(1954). However, Wilson (1954) indicated that some of the mink in

his study were taken in Camden and Dare counties, and no specific

locality was given for the specimen from which the mole was retrieved.

That it was from Currituck County is possibly an erroneous assumption.

It is therefore accompanied by a question mark on the distribution

map. Regardless, this record affirms the presence of Condylura in the

northeastern corner of the state.

Clark et al. (1985) suggested the occurrence of Condylura in

Bladen and Hoke counties (no specific localities given) based on the

presence of mole runs "almost certainly made by this species." Lee

(1987) reported C. cristata from McCain, Hoke County, citing Clark

et al. (1985) as the source. That paper does not, however, contain a

clear reference to the species' occurrence at McCain. Lee (1987)

further reported observation of "burrows and mounds of Condylura"

from West End, Moore County, in 1980. Since these records did not

involve the observation of actual specimens, they are accompanied by

question marks on the distribution map.

Lee (1987) also reported C. cristata from Green [sic] County,

"pre-1950," citing Clark et al. (1985) as the source, but that paper

contains no reference to the species occurring in Greene County. Neither

are there records from that county in the North Carolina State Museum's

files, hence, it is not included on the distribution map.

Two additional reports of C. cristata from the Piedmont of North

Carolina are as follows.

Randolph Co.: 2.1 mi. SSE Ulah, off SR 2843 < 0.1 rdmi. SE

jet. NC 159. John Schneider, a horticulturist with the North Carolina

Zoological Park, reported (personal communication) a specimen that

was captured and badly mangled by his dog in March or April, 1991.

I visited the site in July, 1994, and found no obvious signs of fresh

82 Jeffrey C. Beane

mole activity, but no serious collecting efforts have yet been undertaken

there.

Wake Co.: 1.25 mi. SW Millbrook (= 4.0 mi. N.NE center Raleigh),

4001 Quail Hollow Drive along Big Branch. Dr. Wesley E. Kloos, a

genetics professor at North Carolina State University, reported (personal

communication, NCSM files) having captured two specimens in his

back yard between 1989 and 1991, and released both in nearby Eastgate

City Park. A photograph sent to the North Carolina State Museum, of

Kloos holding one of the captured animals, was too small and blurred

for positive identification. Frequent pitfall trapping on the site during

1992-1993 yielded no specimens.

Although neither Kloos nor Schneider is a mammalogist, both

are scientifically oriented individuals, and each seemed certain of the

animals's identity. The star-nosed mole is certainly a difficult animal

to misidentify. Their reports are therefore included here, and are probably

valid, although they remain unverified, and are accompanied by question

marks on the distribution map. Except for the Surry County record,

and the Mecklenburg County sight record by Brown, they represent

the only reports from the Piedmont region of the state. However, the

records from Polk and Rutherford counties (NCSM 1237 and 6336,

respectively) are very near the eastern edge of the escarpment, and

the sight record of burrows and mounds from West End in Moore

County (Lee 1987) is at the extreme inner edge of the Coastal Plain.

The locality for the Surry County specimen (NCSM 7744) lies

just off the Blue Ridge escarpment in the extreme western Piedmont,

and is the first specimen-supported record for that geographic province

in the state. The specimen was found dead along a bog at the edge of

a hayfield. Evidence of considerable mole activity was apparent at

this site during 1994 and 1995 (personal observation), and the landowners

reported that these moles were commonly killed by house cats on the

property, as was possibly the case with this specimen (Ann B. Somers,

personal communication).

The specimens from Ashe, Brunswick, Dare, Hoke, Jackson, and

Rutherford counties also represent new county records. The sight records

for Bladen, McDowell, Mecklenburg, Randolph, and Wake counties,

though not supported by specimens, represent previously unpublished

county records as well (see Lee et al. 1982, Webster 1987).

DISCUSSION

The range of the star-nosed mole overlaps that of the eastern

mole (Scalopus aquaticus) throughout North Carolina, and that of the

hairy-tailed mole (Parascalops breweri) in the Mountains. The hairy-

Star-nosed Mole 83

tailed mole is currently known from Avery, Buncombe, Caldwell,

Jackson, Haywood, Macon, Mitchell, Swain, Transylvania, Watauga,

Wilkes, and Yancey counties in the Mountains (Lee et al. 1982, NCSM

files). The late Joseph M. Bauman (personal communication) also

reported specimens from Cherokee County. The eastern mole apparently

occurs throughout the state (Lee et al. 1982, Webster et al, 1985),

and is in most places the most common mole.

Differences in habitat preference by the three have been noted

by several authors. Star-nosed moles are known to prefer, if not require,

wet areas as habitat (Hamilton 1931, Burt and Grossenheider 1976,

Yates and Pedersen 1982, Webster et al. 1985). Nearly all North

Carolina specimens for which habitat information is available were

collected in such areas. A possible exception is the Bladen County

specimen listed above, which was found dead in a dry, sandy area

apparently not near a wetland (T. M. Padgett, personal communication).

The eastern mole has been reported to avoid "the wet soils preferred

by the Star-nosed Mole" (Lee et al. 1982). The hairy-tailed mole

reportedly occurs in similar habitats as the eastern mole, but usually

at higher elevations, with most specimens being taken at above 2,000

ft. (610 m) (Lee et al. 1982, Webster et al. 1985, NCSM files). Lee et

al. (1982) reported the hairy-tailed mole to be "considerably more

common than the Star-nosed Mole," but this may not hold true at all

montane localities. In Watauga County, for instance, Condylura apparently

is more frequently encountered than Parascalops (R. W. Van Devender,

personal communication), and there are overall more Condylura than

Parascalops from the North Carolina mountains in the State Museum's

collections.

The ecological niches of the three moles may thus be loosely

described as Parascalops and Scalopus frequenting well-drained soils,

often with an altitudinal separation between them, and Condylura

utilizing wet, low-lying areas at a wide range of elevations. However,

some syntopy may occur. All three species may occur at some montane

localities, as is the case in the vicinity of Highlands Biological Station

in Macon County at ca. 3,900 ft. (1,190 m) in elevation (NCSM

files). In Ashe County at ca. 2,880 ft. (878 m), I found a dead eastern

mole (NCSM 7251) in a wet sedge meadow bog, a site which appeared

more suitable for a star-nosed mole. Eadie (1939) reported having

trapped Condylura and Parascalops in the same tunnel at a New

Hampshire site. Undoubtably, the distributions and interspecific zrelationships

of North Carolina's three mole species merit further study.

The possibility that Condylura cristata occurs at low densities

or at scattered localities throughout the state, and that the Mountain

84 Jeffrey C. Beane

and Coastal Plain populations are not disjunct, should not be ignored.

Although Lee et al. (1982) stated that "absence of records from the

piedmont is certainly not an artifact of collecting," there have probably

been few serious efforts to collect C. cristata in the North Carolina

Piedmont, other than the recent trapping efforts at the Wake County

site. Moreover, specific attempts to collect Condylura in the Mountains

and Coastal Plain have seldom if ever proven successful, and Clark et

al. (1985) commented on the difficulty encountered in trapping the

species. Most available specimens for which the method of collection

is known were found dead on roads or otherwise accidentally encountered.

Even in the Mountains where the species may be fairly common,

there are still several counties for which specimens have not been

reported. Much of the Piedmont, particularly the western part, has

been largely overlooked or ignored by biologists, and its fauna remains

poorly documented. Small, fossorial or otherwise secretive vertebrates

may long elude detection in any region. As examples, the eastern

tiger salamander (Ambystoma tigrinum) was first discovered in the

Piedmont in Wake County by me in 1982 (NCSM files) after that area

had been heavily collected by herpetologists for nearly a century; and

the bog turtle (Clemmys muhlenbergii) was discovered in three new

Piedmont counties during 1992-1993 (Beane 1993). Although some

areas, such as the Raleigh vicinity, have been heavily collected by

mammalogists and other biologists for more than a century, little

effort has been specifically aimed at star-nosed mole trapping.

The North Carolina Piedmont has been heavily altered for agriculture

and urbanization — probably more so overall than either the Mountains

or the Coastal Plain, and many Piedmont wetlands have been drained

or otherwise destroyed in the process. It is possible that wetlands

alteration or other human activities may have eliminated the star-

nosed mole from many areas of the Piedmont in recent times. If

populations of this mole do occur throughout the Piedmont of North

Carolina, it is likely that they exist as scattered relicts (either Pleistocene

relicts or more recent anthropogenic relicts) and at low densities. It is

hoped that this paper will help stimulate biologists working in North

Carolina (and other southern states) to make every reasonable effort

to collect evidence of Condylura, and to photograph or collect specimens

wherever they are encountered, especially at undocumented localities.

ACKNOWLEDGMENTS—^ am grateful to Allen C. Boynton, Elmer

E. Brown, Kurt A. Buhlmann, Dennis W. Herman, Elizabeth McGhee,

Thomas M. Padgett, Philip B. Spivey, R. Wayne Van Devender, W.

David Webster, and David K. Woodward for their assistance in locating

Star-nosed Mole 85

specimens and providing sight records. The discovery of the Surry

County specimen was a result of the Surry County Natural Areas

Inventory, for which Ann Berry Somers and J. Richard Everhart (among

numerous others) deserve special credit. William M. Palmer and two

anonymous reviewers provided helpful comments on earlier drafts of

the manuscript.

LITERATURE CITED

Audubon, J. J., and J. Bachman. 1851. The quadrupeds of North

America. Volume II. New York, New York.

Beane, J. C. 1993. A survey of bog turtle (Clemmys muhlenbergii)

habitat in the western Piedmont of North Carolina. Bulletin of

the Chicago Herpetological Society 28(ll):240-242.

Brimley, C. S. 1923. The star-nosed mole in eastern North Carolina.

Journal of Mammalogy 4(3): 183-184.

Brimley, C. S. 1944-1946. The mammals of North Carolina. 18

installments. Carolina Tips. Carolina Biological Supply Co., Elon

College, North Carolina.

Brown, E. E. 1979. Some snake food records from the Carolinas.

Brimleyana 1:113-124.

Burt, W. H., and R. P. Grossenheider. 1976. A field guide to the

mammals. Third edition. Houghton Mifflin Company, Boston, Massachusetts.

Clark, M. K., D. S. Lee, and J. B. Funderburg, Jr. 1985. The

mammal fauna of Carolina bays, pocosins, and associated com-

munities in North Carolina: An overview. Brimleyana 11:1-38.

Eadie, W. R. 1939. A contribution to the biology of Parascalops

breweri. Journal of Mammalogy 20(2): 150-173.

Hall, E. R. 1981. The mammals of North America. John Wiley and

Sons, New York, New York.

Hamilton, W. J. 1931. Habits of the star-nosed mole, Condylura

cristata. Journal of Mammalogy 12(4):345-355.

Johnston, D. W. 1967. Ecology and distribution of mammals of

Highlands, North Carolina. Journal of the Elisha Mitchell Scien-

tific Society 83(2):88-98.

Lee, D. S. 1987. The star-nosed mole on the Delmarva Peninsula:

zoogeographic and systematic problems of a boreal species in the

South. The Maryland Naturalist 31(2):44-57.

Lee, D. S., J. B. Funderburg, Jr., and M. K. Clark. 1982. A distri-

butional survey of North Carolina mammals. Occasional Papers

of the North Carolina Biological Survey 1982-10. North Carolina

State Museum, Raleigh.

Linzey, D. W., and A. V. Linzey. 1968. Mammals of the Great

Smoky Mountains National Park. Journal of the Elisha Mitchell

Scientific Society 84(3):384-414.

86 Jeffrey C. Beane

Odum, E. P. 1949. Small mammals of the Highlands (North Caro-

lina) Plateau. Journal of Mammalogy 30(2): 179-192.

Paradiso, J. 1959. A new star-nosed mole (Condylura) from the

southeastern United States. Proceedings of the Biological Society

of Washington 72:103-108.

Petersen, K. E., and T. L. Yates. 1980. Condylura cristata. Mamma-

lian Species 129:1-4.

Webster, W. D. 1987. Condylura cristata parva Paradiso, southern

star-nosed mole. Pages 42-43 in Endangered, threatened, and rare

fauna of North Carolina, Part I. A re-evaluation of the mammals

(M. K. Clark, editor). Occasional Papers of the North Carolina

Biological Survey 1987-3. North Carolina State Museum, Raleigh.

Webster, W. D., J. F. Parnell, and W. C. Biggs, Jr. 1985. Mam-

mals of the Carolinas, Virginia, and Maryland. The University of

North Carolina Press, Chapel Hill.

Wilson, K. A. 1954. The role of the mink and otter as muskrat

predators in northeastern North Carolina. The Journal of Wildlife

Management 18(2):199-207.

Yates, T. L., and R. J. Pedersen. 1982. Moles (Talpidae). Pages 37-

51 in Wild mammals of North America. (J. A. Chapman and G.

A. Feldhamer, editors). John Hopkins University Press, Baltimore,

Maryland.

Received 20 October 1994

Accepted 11 April 1995

Observations on North Carolina Crayfishes

(Decapoda: Cambaridae)

John E. Cooper

418 Wayne Drive

Raleigh, North Carolina 27608

AND

Alvin L. Braswell

North Carolina State Museum of Natural Sciences

P.O. Box 29555, Raleigh, North Carolina 27626

ABSTRACT — Cambarus {Tubericambarus) acanthura, Orconectes

(Procericambarus) spinosus, and an apparently undescribed species

of Orconectes (O. sp. B), are reported for the first time from

North Carolina. Six additional specimens of Orconectes (Crockerinus)

virginiensis are reported from the Chowan and Roanoke basins,

and its range in North Carolina is clarified. Orconectes (C.)

erichsonianus, for which a North Carolina locality has been

published, is deleted from the State list. Cambarus {Jugicambarus)

asperimanus is reported from the Watauga and New rivers in

Watauga County, and localities are provided that expand its

known range in the Piedmont Plateau. New locality data, distributional

clarifications, or natural history notes are provided for Cambarus

(Depressicambarus) latimanus, C (D) reduncus, Cambarus {Jugicambarus)

carolinus, C (J) dubius, C. (/.) nodosus, Cambarus (Puncticambarus)

georgiae, C (P.) parrishi, C. (P) reburrus, Procambarus (Ortmannicus)

medialis, P. (O.) pearsei, and P. (O.) plumimanus. A blue

specimen of C. latimanus is reported from the Neuse River

basin. The North Carolina crayfish fauna is correlated with

the State's major river basins and physiographic provinces.

Thanks largely to the efforts of the late Horton H. Hobbs, Jr.,

U. S. National Museum of Natural History, Smithsonian Institution,

with whom the modern era in crayfish studies essentially began, the

composition of the crayfish fauna of North Carolina is relatively well

known. As indicated in his most recent checklist of the American

crayfishes (Hobbs 1989:89), the North Carolina fauna consists of 27

described native species, a putative subspecies of one of them, and

one introduced species. Hobbs, sometimes with coworkers, described

one of the four genera, six of the 11 native subgenera, and 11 of the

27 native species known to occur in the State at the time of his

checklist.

Brimleyana 22:87-132, June 1995 87

88 J. E. Cooper and A. L. Braswell

In addition to the described species, at least four and probably

five known but undescribed species from North Carolina await

description. Hobbs and Peters (1977:8-9) mentioned Cambarus (Cambarus)

sp. A, close to Cambarus (Cambarus) bartonii Fabricius; Cambarus

(Depressicambarus) sp. B, close to Cambarus (Depressicambarus)

reduncus Hobbs; and Cambarus (Puncticambarus) sp. C. The latter,

C. (P.) sp. C, refers not to a single undescribed species but to an

undiagnosed species complex that includes Cambarus (Puncticambarus)

acuminatus Faxon, which is one of the 27 species listed for North

Carolina. In fact, however, Hobbs (1969:135) said that C. (P.)

acuminatus (sensu stricto) may be "confined to the Saluda drainage"

of South Carolina, and later (Hobbs 1989:25) added, "Even in the

Santee Cooper basin (of which the Saluda River is a tributary), . . .

more than one rather distinct 'variant' is recognizable." We see no

reason in our paper to depart from using C. (P.) sp. C for this complex

until a diagnosis has been completed. Considering that the species in

the complex range from the mountains to the coast in North Carolina,

there probably are several awaiting description. Cooper and Cooper

(1977a: 198-199) and Cooper and Ashton (1985:9) commented on the

undescribed Orconectes sp. A, and North Carolina localities for another

apparently undescribed Orconectes (herein designated O. sp. B) are

presented in our paper. One of us (JEC) is investigating a number of

other new species, but further comment on them at this time would

be premature. It is a certainty that other undescribed species await

discovery in North Carolina.

About another widespread crayfish that occurs in parts of

North Carolina, Procambarus (Ortmannicus) acutus acutus (Girard),

Hobbs (1989:64) said, "With little doubt, the populations currently

assigned to this subspecies constitute a species complex." The

complex was under study by Hobbs and Hobbs (1990:608). Hobbs

(1989:24) also said that Cambarus (Lacunicambarus) diogenes Girard,

a broadly distributed crayfish whose range includes eastern North

Carolina, "is a species complex and needs considerable attention."

Jezerinac (1993:532) "concluded that the complex consists of two

subgenera Lacunicambarus and Tubericambarus, new subgenus, and

at least five additional species or subspecies." Another extremely

variable crayfish that occurs in parts of North Carolina, C. (C.)

bartonii, is also in dire need of revision. Hobbs (1969:146) referred

to "the depauperate state of our knowledge of the relationships of

those crayfishes which are currently being tentatively designated as

C. b. bartonii . . . ." Hobbs (1989:82, 89) included Cambarus bartonii

cavatus Hay in his list of crayfishes occurring in North Carolina, but

North Carolina Crayfishes 89

North Carolina was not included in his statement of the range of this

putative subspecies (Hobbs 1989:14). We have opted to omit it as a

member of the North Carolina cambarid fauna until specific evidence

appears to support its inclusion.

Four described species are North Carolina endemics: Cambarus

{Depressicambarus) catagius Hobbs and Perkins, of the upper Cape

Fear River basin; Cambarus (Puncticambarus) reburrus Prins, of the

French Broad and Savannah basins; Procambarus (Ortmannicus)

medialis Hobbs, of the Neuse and Tar-Pamlico basins; and Procambarus

{Ortmannicus) plumimanus Hobbs and Walton, of the Northeast

Cape Fear and New (White Oak) basins, which, as we explain later,

may be expanding its range into the lower Neuse basin. The undescribed

Orconectes sp. A is another North Carolina endemic, found only in

the Neuse and Tar-Pamlico basins.

This article adds two described and one undescribed native species

to the State list, provides additional distributional and natural history

information for them and 13 other species, deletes a species for which

a North Carolina locality has been published, correlates the State's

crayfish fauna with its major hydrologic units, and summarizes the

distribution of this fauna in the State's three major physiographic

provinces.

Abbreviations used are as follows. SR = secondary road (formerly

CR for county road); NC = North Carolina state highway; US = United

States highway (an A after the number means alternate highway); I =

interstate highway; FRS = United States Forest Service Road; cntr =

center of town or city; NCSM C- = crustacean collections, North

Carolina State Museum of Natural Sciences, Raleigh; USNM = collections,

U. S. National Museum of Natural History, Smithsonian Institution,

Washington, D. C. Collector's names are given in first usage, initials

thereafter.

Cambarus {Depressicambarus) latimanus (LeConte)

Hobbs (1981:120), reporting observations made in a thesis by

J. L. Boyce (1969), stated that in Yellow River, Georgia, C. latimanus

"becomes relatively inactive during December, January, and

February," Hobbs (1981:119) also provided monthly capture data for

a large number of specimens that suggested the same thing: 77 collected

in November, five in December, 21 in January, and none in February,

as opposed to 164 in March, 1,079 in April, and relatively

high numbers in most other months. These kinds of collecting data,

however, can be more a reflection of seasonal activities of the collectors

than of the collected. If we can equate "activity" with "catchability,"

90 J. E. Cooper and A. L. Braswell

and in the case of animals taken in traps we feel justified in so

doing, C. latimanus is not inactive during winter months in eastern

North Carolina. Intensive sampling was conducted in the Neuse River

basin from late December 1978 through May 1979, and in the Tar-

Pamlico basin from January through April 1980, most of it part of a

study of the distribution and ecology of the salamander, Necturus

lewisi (Brimley) (Braswell and Ashton 1985). This effort yielded

considerable winter and spring capture ("activity") data for 409 adult

and 279 juvenile C. latimanus, the majority of them caught in

traps (Table 1).

Bouchard (1978:37) remarked on a dearth of ovigerous females

in collections of this species, saying, "Of the numerous collections of

C. latimanus that have been made, only three specimens are females

with eggs (or young)." One of these females was collected in a

tributary of the Neuse River in Wake County, North Carolina, on 16

April 1977, the other two were collected in Alabama on 22 April

1973. Hobbs (1981:119), who examined 2,424 specimens of this species

from approximately 400 localities, remarked, "I have no records of

ovigerous females or of those carrying young anywhere within its

range .... This paucity of such females in collections almost certainly

reflects inadequate sampling of burrows in the banks of streams."

Most of the smallest juveniles collected in the Neuse and Tar-Pamlico

surveys were taken in late April and May. This, combined with the

few available dates of capture of ovigerous females, Bouchard's (1978:47)

report of a copulating pair found on 2 November 1974 in Alabama,

and the fact that nearly 50 percent of the males in the January sample

reported in Table 1 were form I, leads us to suggest that in eastern

North Carolina (1) mating and egg production occur in winter and

perhaps late fall, (2) incubation occurs during winter and early spring,

Table 1. Summary of winter and spring captures of C. latimanus in

the Neuse and Tar-Pamlico river basins, 1978-1980.

North Carolina Crayfishes 91

and (3) the young are produced later in the spring (a time of high

water, expanded habitat, and increased availability of food). This

supports Thorp's (1978:278) statement that in the lower Savannah

River basin of South Carolina this species "enters the reproductive

period as temperatures are dropping in fall and winter."

On 17 February 1982, a blue specimen of C. latimanus was

found by a worker clearing a muddy ditch at the south end of the

Wayne Community College campus in Goldsboro, Wayne County

(Neuse River basin). It was taken to Gary W. Woodyard of the

college biology faculty, who generously donated it to the N. C. State

Museum. The animal, a form II male (NCSM C-775), was generally

cobalt blue, with the pigment obviously in the exoskeleton since the

underlying chromatophore pattern of blotches and abdominal

stripes was clearly visible. The right cheliped was missing, but the

left cheliped and all pereiopods were whitish on the ventral surfaces,

with a pale bluish tint at the base of the coxa of each limb. The

lateral margin of the palm and propodus was very light blue, but

the entire dactyl was darker except at the ventral tip. The gonopods

(first pleopods), eyestalks, antennae, antennules, and antennal scales

were pale blue. There was a small white area on the anterolateral

carapace just below the suborbital angle and along part of the margin

of the carapace. The small tubercles and punctations of the carapace

were points of white, as were the small cervical spines. Most of

the ventral abdomen was clear, but the transverse ridges between

segments were blue.

This is the first blue individual reported for this species, and

the first blue crayfish reported for any non-blue species in North

Carolina. One North Carolina crayfish, Cambarus (Jugicambarus)

dubius Faxon, is known to have a cobalt blue color morph. In the area

where the blue C. latimanus was found, the species normally is light

tan or greenish, with dark brown or green carapace markings and

abdominal stripes. Hundreds of normal-colored specimens of this

species have been collected in the Neuse River and its tributaries,

including a number from Wayne County.

Fitzpatrick (1987) summarized most of the known records for

the "blue color phase" in six crayfish genera, and discussed environ-

mental (diet and illumination) versus genetic causes of such color

variations. Penn (1951) reported a blue color variant of Procambarus

(Scapulicambarus) clarkii (Girard), a normally red species, in

Louisiana. Penn later (1959:10) said, "In recent years anomalous bright

blue specimens have appeared in several parts of the state . . . ."

(That someone may be culturing this obviously genetic variant is

92 J. E. Cooper and A. L. Braswell

indicated by the fact that in April 1984 JEC examined several bright

blue specimens of this species, said to be from "Thailand," that

were for sale as exotic novelties in a Raleigh tropical fish shop. A

form I male was purchased and is in the NCSM collections, C-1295.)

Smiley and Miller (1971:221) estimated the frequency of blue

variants in normally non-blue P. a. acutus as 1 in 50,000. Black

(1975) experimentally demonstrated that blue color in this species is

a mutation, in which the gene controlling the chemical composition

of a carotenoid pigment in the exoskeleton behaves as a single

recessive allele, with complete penetrance of the dominant normal

allele. He estimated the ratio of blue to normal crayfish in the pond

of origin of his parental stocks at 1:5,600. Anthony D'Agostino has

bred cobalt blue Homarus americanus, and the Fx offspring inherit

this coloration as a homozygous recessive trait (Porterfield 1982:38),

which is what Black found in P. a. acutus.

With a nod to William of Occam, we find it considerably more

parsimonious to conclude that the Wayne County C. latimanus was a

genetically blue individual than that its abnormal color was produced

by diet or environmental conditions.

In North Carolina, C. latimanus is a common, widespread

inhabitant of the eastern Piedmont Plateau and much of the Coastal

Plain. It is possible, though, that it also occurs in the Hiwassee River

basin of the Blue Ridge, as we discuss later.

Cambarus (Depressicambarus) reduncus Hobbs

Bouchard (1978:40) gave the range of this species as "in the

Piedmont province from the Cape Fear River drainage in North

Carolina southward to the Santee River basin . . . .," and in the latest

American checklist Hobbs (1989:16) gave it as "Piedmont Province

from Orange County, North Carolina to Richland County, South

Carolina." Concerning the northern terminus of its range, these

statements almost certainly were based on specimens from the upper

Cape Fear basin in and around Chapel Hill, Orange County, earlier

reported by Hobbs (1956:66-67) and Hobbs and Peters (1977:50).

Although Cooper and Ashton (1985:9) had reported C. reduncus from

the Neuse and Tar-Pamlico basins, north of the Cape Fear, they gave

no particulars. The following localities extend the known range of

this species north into the Piedmont headwaters of these two rivers,

very close to the Roanoke River basin.

NEUSE RIVER BASIN. Durham Co.— (1) Lick Crk at SR 1905,

10 air mi (16 air km) E Durham; 1 6 I (NCSM C-520), 27 Jan 1979,

A. P. Capparella. Granville Co. — (2) burrow in roadside ditch along

North Carolina Crayfishes 93

SR 1721, 3.5 air mi (5.6 air km) SE cntr Creedmoor, near Wake Co

line; 1 6 I (NCSM C-860), 23 Apr 1978, R. E. Ashton, Jr.; (3)

swamp at E edge Lake Rogers (on Ledge Crk), 1 air mi (1.6 air km)

NW Creedmoor; 1 j ^ 9 Mar 1991, D. G. Cooper, JEC. Wake Co.—

(4) burrow in lawn, Morrisville; 1 9 (NCSM C-907), 27 Mar 1978, J.

Clayton; (5) alive on SR 1300 (Hemlock Bluffs Rd) N of bridge; 1 6

I (NCSM C-909), night 3 Nov 1977, R. W. Laney, D. F. Lockwood;

(6) alive on road, 0.5 mi (0.8 km) W jet SR 1379 & Kildaire Rd; 1 6

I (NCSM C-910), night 3 Nov 1977, RWL, DFL. TAR-PAMLICO

RIVER BASIN. Granville Co.— (7) alive on road, jet SR 1304 &

1307, 1.8 air mi (2.9 air km) W Hebron; 1 6 II (NCSM C-1262),

night 23 Apr 1983, J. P. Kumyhr, D. Smith. Person Co.— (8) Tar R at

SR 1565, 1.7 air mi (2.7 air km) WSW Dennys Store; 1 6 I, I 9

(NCSM C-665), 20 Feb 1980, trap, E. Rawls. Vance Co.— (9) Tabbs

Crk at SR 1101, 2.3 air mi (3.7 air km) W Kittrell; 1 6 I, 1 9

(NCSM C-655), 24 Jan 1980, trap, ER.

Hobbs and Peters (1977:18) reported C. reduncus from "Mont-

gomery County: (2) Hamer Creek, 3 mi (4.8 km) N Richmond Co line

on St Rte 73," Assigning the locality to the "CATAWBA BASIN."

This locality, though, like nearly all of Montgomery County, is in

the Yadkin-Pee Dee basin. The following collections of C. reduncus,

however, are known from the Catawba basin in North Carolina:

Gaston Co. — (1) small stream (trib South Crowders Crk) near office

Crowders Mountain State Park, off SR 1125, 0.6 rd mi (0.9 rd km)

SW jet SR 1106, 5.2 air mi (8.3 air km) S Bessemer City; 1 6 II, 1

j 6, 3 9, 4 j 9 (NCSM C-2300), 22 Apr 1985, ALB. Mecklenburg

Co.— (2) plowed field along dirt rd at E end SR 3629 (Sixmile Crk

dr), ca. 6.3 air mi (10.1 air km) SE cntr Pineville; 1 6 II, 4 j 8, 1 9 ,

2j 9 (NCSM C-1043), 19 Apr 1980, from burrows with chimneys,

N. L. Elliott; (3) ditch along SR 3629 (Sixmile Crk dr), ca. 5.6 air mi

(8.9 air km) SE cntr Pineville; 9 j 8, 1 9, 11 j 9 (NCSM C-1045),

19 Apr 1980, NLE. Union Co.— (4) along SR 1624, 0.5 rd mi (0.8 rd

km) NW jet NC 200, 2.0 air mi (3.2 air km) NNE Monroe; 1 8 I

(NCSM C-311), 15 Jul 1978, J. W. Braswell, Jr., ALB; (5) East Fk

Twelve Mile Crk at SR 1329, ca. 2.1 air mi (3.4 air km) SSW Wesley

Chapel; 1 j 8, 3 j 9 (NCSM C-1031), 18 Apr 1980, in roadside ditch

fed by woodland pool; 1 j 8, 3 j 9 (NCSM C-1047), 18 Apr 1980, in

shallow, temporary floodplain pools; 1 8 II, 1 j 9 (NCSM C-1027),

26 Apr 1980, dug from burrows with chimneys in old-field, NLE; (6)

small branch Little Twelve Mile Crk at SR 1329, ca. 2.9 air mi (4.6

air km) SSW Wesley Chapel; 4 j 8, 7 j 9 (NCSM C-1037), 18 Apr

1980, NLE; (7) woodland pool off Little Twelve Mile Crk at SR 1328,

94 J. E. Cooper and A. L. Braswell

ca. 3.1 air mi (5.0 air km) SW Wesley Chapel; 3 j 8, 1 j 9 (NCSM

C-1040), 18 Apr 1980, NLE; (8) damp ditch at East Fk Twelve Mile

Crk at SR 1336, ca. 1.6 air mi (2.6 air km) SW Wesley Chapel; 1 j 8,

1 j 9 (NCSM C-1038), 20 Apr 1980, from burrows, NLE.

Cambarus (Jugicambarus) asperimanus Faxon

This crayfish occurs in the Watauga River basin in Tennessee

(Hobbs 1989:20), but no localities for the species have been reported

from this system in North Carolina. Additionally, no one unequivocally

has reported its occurrence anywhere in the New River basin. With

respect to the latter, in a discussion of "Cambarus (Cambarus)

bartoni asperimanus" Ortmann (1931:138) stated, " I have a male (I)

and two females from Blowing Rock (in the headwaters of New River,

some distance northeast of Asheville), in which the inner

margin of the palm has the tubercles somewhat more strongly develop-

ed than is usual in bartoni. This indicates a transition toward

asperimanus. However, I prefer to leave these with bartoni; the

character being much less distinct here than in the specimens from

Asheville and Canton" (Buncombe and Haywood counties, respectively,

French Broad River basin). There is one other published locality for

C. asperimanus that could be in the New River basin, but it was not

reported as such. Hobbs and Peters (1977:57) recorded the species

from "Watauga County: (8) creek at Deep Gap," but placed this locality

in the Pee Dee (Yadkin) basin. This citation does not indicate whether

"creek at Deep Gap" refers to the community of Deep Gap, or the

mountain gap of the same name. The community is in eastern Watauga

County on Deep Gap Creek, a tributary that joins South Fork New

River in Ashe County. The mountain gap lies southeast of the Blue

Ridge, in the drainage of Stony Fork Creek, a headwater tributary

that enters the Yadkin River in western Wilkes County.

On 25 July 1984, we collected C. asperimanus in both the

Watauga and New River basins at the following localities. WATAUGA

RIVER BASIN. Watauga Co.— (1) small stream entering N bank

Watauga R at NC 105 bridge, jet SR 1112, ca. 2.4 air mi (3.8 air km)

SE Valle Crucis; 2 $ (NCSM C-1811), from under separate rocks at

mouth of stream. NEW RIVER BASIN. Watauga Co.— (2) small,

shallow creek (South Fk New River dr) in hardwood ravine and

meadow on E side Howard Knob, off Hidden Valley Circle, 0.3 rd mi

(0.5 rd km) from jet Chestnut Drive in north Boone; 1 cM, 5 j 8,1 9,

1 j 9 (NCSM C-1814), 1 9 with attached young (NCSM C-1815),

3 9, 1 j 9 (NCSM C-1816), with R. W. VanDevender.

North Carolina Crayfishes 95

At the Watauga River locality, the only other crayfish collected

was Cambarus (Puncticambarus) robustus Girard, all specimens of

which were taken from the river itself. At the New River locality, the

orangish-tan C. asperimanus were "associated" with cobalt blue C.

dubius. The two species were dug from burrows within a few meters

of each other, but their habitats differed. The C. asperimanus that

came from burrows were in simple, shallow burrows at or just above

water level along the banks of the creek, while the C. dubius were

dug from more complex burrows in a mucky seepage area near the

same creek. Some C. asperimanus were found under rocks in the

creek, but no C. dubius were in this habitat, and no C. asperimanus

were dug from the seep.

The limits of the range of C. asperimanus east of the Blue Ridge

escarpment in North Carolina are not yet clear, but we made a number

of collections of the species at the base of the eastern foothills and

in the Piedmont as far east as western Catawba County. Hobbs and

Peters (1977:57) first recorded the species in this area, from a locality

in the upper Catawba basin in northwestern Burke County. Our

collections at the following new localities extend the range of C.

asperimanus farther east in the Catawba basin, and south and east

into the Broad River basin.

CATAWBA RIVER BASIN. Burke Co.— (1) Jacob Fk nr office

South Mountain State Park, 3.4 air mi (5.4 air km) SW Pleasant

Grove; 1 S I, 3 S II, 1 j S, 4 9 (NCSM C-2281), 23 Apr 1985; (2)

Laurel Crk along NC 18, 2.0 rd mi (3.2 rd km) ESE jet SR 1929, 3.2

air mi (5.1 air km) N Pleasant Grove; 2 S I, 5 jS, 6 j 9 (NCSM C-

2286), 1 ovig 9 (NCSM C-2309), 26 Apr 1985. Catawba Co.— (3)

small stream (trib Henry Fk) along I 40 at milepost 120 near Burke

Co line, ca. 4.0 air mi (6.4 air km) SW cntr Hickory; IS II, 3 j S, 1

9, 1 j 9 (NCSM C-1257), 12 Apr 1983. McDowell Co.— (4) trib

Swannanoa Crk along SR 1400, 1.3 rd mi (2.1 rd km) W jet E end

SR 1407, 2.1 air mi (3.4 air km) WNW cntr Old Fort; 2 S I, 2 S II, 1

j 6 (NCSM C-807), 20 Aug 1977; (5) trib Catawba R along US 70,

3.1 mi (5.0 km) E Buncombe Co line, 2.3 air mi (3.7 air km) WSW

cntr Old Fort; 2 9 (NCSM C-949), 24 Nov 1978, with D. L. Stephan;

(6) small cascading stream (trib Swannanoa Crk) along SR 1407, ca.

0.4 rd mi (0.6 rd km) NE Buncombe Co line, ca. 5.0 air mi (8.0 air

km) W cntr Old Fort; 1 j 6,3 9, 1 j 9 (NCSM C-1251), 12 Apr

1983; (7) Long Br (trib Mill Crk) along SR 1407, 0.4 rd mi (0.6 rd

km) S railroad crossing, 1.5 air mi (2.4 air km) SE Graphite & 3.2

air mi (5.1 air km) WNW cntr Old Fort; IS II, 1 j 9 (NCSM

C-1258), 12 Apr 1983; (8) Buck Crk at NC 80 bridge, 0.4 rd mi

96 J. E. Cooper and A. L. Braswell

(0.6 rd km) N jet SR 1437, ca. 0.2 air mi (0.3 air km) E Sunny Vale;

1 9 (NCSM C-2200), 26 Sep 1984. BROAD RIVER BASIN. Cleveland

Co. — (9) small stream (Buffalo Crk dr) in hardwoods, 0.8 rd mi

(1.3 rd km) S Lincoln Co line, 2.3 air mi (3.7 air km) NNE Belwood;

1 8 II, 3 j 8, 3 j 9 (NCSM C-2283), 22 Apr 1985. McDowell Co.—

(10) intermittent creek (trib Cane Crk) in hardwood ravine along

NC 226, 0.1 rd mi (0.2 rd km) WNW Rutherford Co line, 3.4 air mi

(5.4 air km) SE Dysartville; 4 8 II (NCSM C-987), 25 Jun 1977, 2 8

I, 5 8 II, 11 j 8, 9 9, 14 j 9 (NCSM C-1821), 1 9 with attached

young (NCSM C-1823), 1 8 II, 1 j8, 2 $ (NCSM C-1824), 26 Jul

1984. Rutherford Co.— (11) trib Broad R at NC 9, ca. 0.3 air mi

(0.5 air km) S town Lake Lure; 1 8 II (NCSM C-64), 6 Aug 1976;

(12) small stream in steep ravine at NC 9, 0.8 rd mi (1.3 rd km) W

cntr town Chimney Rock; 1 9 (NCSM C-959), 3 8 II, 1 9 (NCSM

C-960), 5 Jun 1977, with E. Messersmith. Polk Co.— (13) unnamed

trib Green R near Raccoon Mountain; 1 8 I (NCSM C-2012), summer

1974, G. G. Shaw.

We conclude, based on the apparently broad temporal distribution

of ovigerous females and those with young, that C. asperimanus

either has an extended breeding season or has a long development

time for the ova and young. Hobbs (1981:190) reported an ovigerous

female taken in Rabun County, Georgia, in April 1977, and said that

"elsewhere" such females have been taken in April, June, and

December. The latter statement may have been based on Bouchard

(1972:47), who reported ovigerous females (presumably in Tennessee)

in "April to June and December." As indicated above, we collected

a female with attached young in Watauga County on 25 July 1984,

and another a day later in McDowell County. David G. Cooper collected

a female with three attached young in Jackson County on 10 August

1993. In addition, we found a number of individual females

closely associated with groups of tiny young in Macon County on 19

September 1984, and on this same date we found free-living young of

the same size.

The Jackson County female collected by DGC on 10 August

1993 is the largest C. asperimanus we have seen, with a total carapace

length (TCL) of 42.0 mm and a postorbital carapace length (PCL) of

37.0 mm. Another very large specimen, a form I male measuring 38.0

mm TCL (32.5 mm PCL), was collected by DGC at the same site on

the same date. We collected a form I male in Macon County that

measured 36.0 mm TCL (31.5 mm PCL). The only other size data we

know of for this species are those of Hobbs (1981:190). His largest

North Carolina Crayfishes 97

Georgia specimen, an ovigerous female, measured 30.5 mm TCL (26.7

mm PCL), and his largest form I male was 27.6 mm TCL (24.1 mm

PCL).

Cambarus (Jugicambarus) carolinus (Erichson)

Although Hobbs and Bouchard (1973:42), Bouchard (1976:594),

and Hobbs (1989:21) included southwestern North Carolina, south

and west of the French Broad River, within the range of this species,

no one has yet reported precise locality or natural history data for

this primary burrower in North Carolina. Faxon (1890:624) reported a

form I male of his Cambarus dubius, collected by James Mooney

from " 'Among the Cherokees,' Indian Territory," and said, "According

to the label accompanying the specimen it is called Tsisgdgili (red

crayfish) by the Cherokee Indians." The same author later (Faxon

1914:396) included this specimen (USNM 14314) under C. carolinus,

said "I am advised by Mr. Mooney that it was in reality obtained

in Swain Co. or in Jackson Co., N. C, " and added (Faxon 1914:397)

that "The living color . . . was red . . . ." Ortmann (1931:147) reported

Faxon's specimen under "Cambarus (Cambarus) carolinus ," and

inexplicably indicated that it was from "Cherokee Co.; North Carolina."

He added, "I have seen, at Murphy, Cherokee Co., crawfish-chimneys,

and tried unsuccessfully to dig out specimens .... The owner of the

place told me, that these were red crawfish, and thus probably this

species." Ortmann (1931:147) also recorded the species from "Swampy

ground near springs, Marion, McDowell Co. . . . Swamp, Ashville

[sic], Buncombe Co., . . .," and "Blowing Rock, Watauga Co., . . . ."

These three localities are outside the range of C. carolinus as

currently understood, but the last two are within the known range of

C. dubius (sensu lato). The McDowell County locality is in the western

Piedmont Plateau, in the headwaters of the Catawba River basin,

beyond the known ranges of both C. carolinus and C. dubius. Brimley

(1938) recorded C. carolinus at "Judson," and Hobbs and Bouchard

(1973:21) called for confirmation of this record. This would have

been a reasonable place to find C. carolinus, since Judson was a

community on the Little Tennessee River in southwestern Swain

County, but it is now beneath the waters of Fontana Lake (Powell

1968:259). The reference by Harris (1903:142) to "C. carolinus" in

the Tar-Pamlico River basin was clearly in error.

We made the following collections of specimens referred to

this species at the indicated localities, all within the basin of the

Little Tennessee River: Clay Co. — (1) boggy spring seep in Riley

Cove, off unnumbered dirt road reached from US 64, 3.3 rd mi (5.3 rd

98 J. E. Cooper and A. L. Braswell

km) S jet SR 1359 & 1.2 rd mi (1.9 rd km) W jet FSR 71 (old US 64),

Natahala National Forest; 1 8 II (NCSM C-948), 16 Aug 1977, 1 8

I, 1 8 II, 5 j 8, 4 9, 3 j 9 (NCSM C-2165), 1 <? II, 1 9 (NCSM

C-2166), 21 Sep 1984. Graham Co.— (2) dug from muck in Talulah

(Tulula) Bog, along US 129, 1.4 rd mi (2.2 rd km) E jet SR 1201, 6.7

air mi (10.7 air km) SE Robbinsville; 1 j 8,2 9, 1 j 9 (NCSM

C-2290), 25 Apr 1985; (3) small boggy area near Cheoah R along

SR 1147, 0.1 rd mi (0.2 rd km) S jet US 129; 1 ovig 9, 1 j 9 (NCSM

C-2310), 26 Apr 1985. Macon Co. — (4) boggy area around flood-

plain pond by Nantahala R, off US 19, ca. 0.5 rd mi (0.8 rd km) NE

Graham Co line & 5.2 air mi (8.3 air km) NW Kyle; 1 8 II, 1 j 8, 1

9 , 1 j 9 , remains of decomposed 8 I, pair of loose chelipeds (NCSM

C-2292), 24 Apr 1985, with J. Bauman; (5) dug from sphagnum

bog and margin murky stream along Little Choga Crk on SR 1402,

3.3 rd mi (5.3 rd km) SE jet SR 1401, 3.2 air mi (5.1 air km) WSW

Aquone; 1 8 II, 1 9 (NCSM C-2287), 25 Apr 1975, with JB;

(6) under rocks in wet roadside ditch along SR 1401, 2.9 rd mi (4.6 rd

km) NE jet SR 1402, 2.3 air mi (3.7 air km) WNW Aquone; 2 j 8

(NCSM C-2289), 25 Apr 1985, with JB.

All the specimens were brick red in color, and all except those

from locality (6) were dug from burrows, usually constructed among

soil and roots in boggy areas. The bog at locality (1) was about 8 m

from a small, rocky stream, tributary to Buck Creek (Nantahala River).

Only one other crayfish, a juvenile male C. b. bartonii, was dug from

this bog. In the nearby stream, under rocks and in substrate under

rocks, we found other C. b. bartonii, and a female of an unidentified

Cambarus. The stream was about 1- to 1.5-m wide, 7- to 8-cm deep,

and had a low gradient and low velocity current. At locality (4), an

unidentified Cambarus was found under a rock in wet sand near the

bog, and at locality (5) C. b. bartonii was found in the stream but

not in the boggy area.

Cambarus (Jugicambarus) dubius Faxon

There are a number of published localities for C. dubius in North

Carolina, including recent ones in Hobbs and Peters (1977:24, 50;

1989:324). The range of the species in North Carolina, however, is

poorly known, both because it is a primary burrower and difficult to

find and collect, and because of taxonomic uncertainties. In general,

the species occurs in "northwestern North Carolina" (Hobbs 1989:

22), "north and west of the French Broad River basin" (Bouchard

1976:594). Little has been published about its natural history in North

Carolina.

North Carolina Crayfishes 99

The following collections of C. dubius are of distributional

and natural history interest. NEW RIVER BASIN. Watauga Co.—

(1) seepage area near small, shallow creek (South Fk New R dr) in

hardwood ravine and meadow on E side Howard Knob, off Hidden

Valley Circle, 0.3 rd mi (0.5 rd km) from jet Chestnut Drive, in north

Boone; 3<?II, 8jd,2$,8j$ (NCSM C-1812), 2 9 with attached

young (NCSM C-1813), 2 j 9 (NCSM C-1817), 1 6 I, 5 6 II, 3 j 6,

2 9, 5 j 9 (NCSM C-1818), 25 Jul 1984, RWV, ALB, JEC. FRENCH

BROAD RIVER BASIN. Haywood Co.— {2) bog near small stream

(trib Cold Springs Crk, Pigeon R dr), in rhododendron thicket in steep

ravine along FSR 148, Pisgah National Forest, 3.6 rd mi (5.8 rd km)

SW jet SR 1334; 1 6 I, 2 6 II, 4 j 9 (NCSM C-2192), 23 Sep 1984,

ALB, JEC. YADKIN-PEE DEE RIVER BASIN. Surry Co.— (3) Home

Crk Off SR 2072, ca. 8.3 air mi (13.3 air km) S town Pilot Mountain;

1 S II (NCSM C-95), 20 Jul 1976, R. M. Shelley, ALB; (4) open

bog Schuyler pasture, ca. 0.6 mi (1.0 km) N Low Gap, between

E side NC 89 and Gulley Crk (Fisher R dr); 1 9, 19 Aug 1994, A.

B. Somers. Wilkes Co. — (5) Stone Mountain State Park; 1 6 I

(NCSM C-1106), 16 Jul 1975, D. S. Lee, P. Hertl; (6) Hunting Crk

at SR 2428, 0.3 rd mi (0.5 rd km) S NC 115, ca. 9 air mi (14.4 air

km) SE Wilkesboro; 1 9 (NCSM C-1234), 15 Jul 1976, F. D. Scott,

M. E. Filka.

At the Watauga County site the C. dubius, cobalt blue in

color, were dug from burrows in a mucky seep near the creek. One of

the females (NCSM C-1813) had young attached to its pleopods.

Thirteen specimens of C. asperimanus also were collected at this

site, but occupied a different habitat than the C. dubius (see the C.

asperimanus account). At the Haywood County site the C. dubius,

this time of a brick red color morph, also were dug from burrows in a

boggy area, about 3 to 4 m from a small stream. The form I male, one

of the form II males, and two of the juvenile females showed consider-

able exoskeleton decalcification, and the other form II male was soft.

In the stream near this bog, eight C. b. bartonii were found under

rocks or dug from the stream substrate. The female from site (4)

in Surry County was found walking in wet grass at about 1900

hours under rainy conditions.

Cambarus (Jugicambarus) nodosus Bouchard and Hobbs

Only two specimens of this burrowing crayfish, females from

two separate localities in Cherokee County, have been reported from

North Carolina (Bouchard and Hobbs 1976:13). We collected 45 specimens

of this species from two additional localities in the Hiwassee River

100 J. E. Cooper and A. L. Braswell

basin: Cherokee Co. — (1) seepage area on slope above SR 1322 and

Shuler Crk, 0.1 rd mi (0.2 rd km) E of Tennessee state line, ca. 4.3

air mi (6.9 air km) WSW Violet; 4 6 II, 12 j 6,2 9, 8 j 9, 1 j

unsexed (NCSM C-1786), 1 6 I, 1 9 (NCSM C-1787), 21 Jul 1984;

(2) seeps and shallow intermittent water on hillside above SR 1323

and trib Shuler Crk, 2.8 rd mi (4.5 rd km) W jet SR 1324, ca. 3.3 air

mi (5.3 air km) WSW Violet; l<JI,4<?n,5jd,39,3j9 (NCSM

C-1788), 21 Jul 1984.

All specimens were dug or scraped from shallow burrows in

dark muck. Some of the burrows were submerged in seepage water,

and some specimens were found under rocks in these areas. No other

crayfishes were found with C. nodosus.

In North Carolina, C. nodosus is limited to the Hiwassee River

basin, where it appears to be the ecological equivalent of C.

asperimanus. The two species have never been reported from the

same localities or even the same river basins anywhere, and C.

asperimanus, although common and widespread in other montane

(and western Piedmont) river systems, seems to be absent from the

Hiwassee River and its tributaries.

Cambarus (Tubericambarus) acanthura Hobbs

Although known from two localities in Fannin County, Georgia

(Hobbs 1981:219-220), this crayfish has never been reported

from North Carolina. Three localities in the Hiwassee River basin

of North Carolina are now known: Cherokee Co. — (1) floodplain

pond and ditches along Nottely R off end SR 1404, ca. 1.0 mi

(1.6 km) N US 64 and 5.8 air mi (8.0 air km) SW Murphy; 1 6 I, 2

j 9, 1 carapace (NCSM C-529), 24 Nov 1978, DLS ALB, 5 j 6\ 2

j 9 (NCSM C-2291), 24 Apr 1985, JB, ALB; (2) floodplain

Nottely R at US 64 Bridge, 0.8 mi (1.3 km) E Ranger; 5 j 8, 3 j

9 (NCSM C-496), 1 6 II (NCSM C-497), Z j 6 (NCSM C-498), 1

6 I (NCSM C-499), 17 Dec 1976, FDS, ALB; (3) small creek

(trib Nottely R) in rhododendron thicket off SR 1423, ca. 0.5 mi (0.8

km) N US 64, 5.0 air mi (8.0 air km) SW Murphy; 1 j 9 (NCSM

C-1790), 22 Jul 1984, ALB, JEC.

At locality (3), the only other crayfish found in the creek was

a male II Cambarus (Puncticambarus) hiwasseensis Hobbs. Speci-

mens C-497 and C-499 were kept alive in the laboratory after their

capture in December 1976; the former died on 13 May 1980, and the

latter, which was a juvenile when collected, died a form I male on 22

May 1981. The pond at locality (1) had extensive mats of floating

North Carolina Crayfishes 101

water shield (Brasenia sp.) when we visited it on 21-22 July 1984. A

trap set there overnight yielded no crayfish.

Cambarus (Puncticambarus) georgiae Hobbs

This crayfish has been reported from only three localities, one

in Rabun County, Georgia (the type locality), and two in Macon

County, North Carolina, all in the upper Little Tennessee River basin

(Hobbs 1981:255). The second North Carolina locality was given as

"Sugar Fork River 8 mi NE of Franklin on US Hwy 64 ... 26 June

1957, E. A. Crawford." In response to a query about this site, Hobbs

(in litt.) said it should be emended to Cullasaja River, US 64 at

Gneiss, 8 mi (12.8 km) SE of Franklin. According to Powell (1968:130),

"The Cherokee work, Kul-say-gee, means 'sugar' or 'sweet,' " and at

least one part of the river apparently was known in the 19th century

as Sugar Town Creek.

We collected 39 C. georgiae at three localities in the Cullasaja

River watershed: Macon Co. — (1) Buck Crk at culvert under US

64-NC 28, 0.2 rd mi (0.3 rd km) S jet SR 1535, ca. 0.6 air mi (1.0 air

km) S Gneiss; 1 6 I, 2 9, 2 j 9 (NCSM C-2155), 20 Sep 1984; (2)

Cullasaja R at bridge, jet SR 1667 & 1653, ca. 1.0 air mi (1.6 air km)

SSE airport in Franklin; 1 j 6 (NCSM C-2161), 20 Sep 1984;

(3) Cullasaja R along US 64-NC 28, 0.2 rd mi (0.3 rd km) N jet

SR 1678, ca. 0.2 air mi (0.3 air km) S Gneiss; 6 j 6,9 j 9 (NCSM

C-2158), 3 6 I, 1 6 II, 1 j 6, 5 9 (NCSM C-2159), 4 j 6, 4 j 9

(NCSM C-2167), 21 Sep 1984; this locality is either the same as,

or very close to, the second North Carolina locality provided by

Hobbs (1981:255).

At locality (1), a single female C. asperimanus and 22 C. b.

bartonii were also taken. At site (3), we also found 29 C. b. bartonii.

On 19 and 20 September 1984, we made collections at three

localities in the Cullasaja drainage above Cullasaja Falls, at elevations

near or above 900 m (3,000 ft). At the highest of these sites we found

only a large number of C. asperimanus, and at the others we found

only 46 C. asperimanus and 51 C. b. bartonii. Thus, C. georgiae

almost certainty is absent from higher elevation, high-gradient streams

in the Cullasaja watershed.

We can add a second Georgia locality to the range of the

species: Rabun Co. — Little Tennessee R at Hwy 246, ca. 0.3 mi (0.5

km) E jet US 441, NNE of Dillard; 3 6 I, 1 j 6 (NCSM C-626), 20

Oct 1979, REA, Jr, J. Perry.

102 J. E. Cooper and A. L. Braswell

Cambarus (Puncticambarus) parrishi Hobbs

This species, which is endemic to the Hiwassee River basin,

previously was known in North Carolina from two collections made

in Clay County in 1959 and 1960 (Hobbs 1981:267). On 22

September 1984, we collected 21 specimens at two additional

sites: Clay Co.— (1) Fires Crk along FSR 340, 2.3 rd mi (3.7 rd km)

NE of end SR 1344 (which becomes FSR 340), Nantahala National

Forest, ca. 4.3 air mi (6.9 air km) NNW cntr Hayesville; 1 6 I, 2 9,

1 j 9 (NCSM C-2174); (2) trib Fires Crk (likely Rockhouse Crk)

along FSR 340A, ca. 0.8 rd mi (1.3 rd km) N jet FSR 340 and 2.9

rd mi (4.6 rd km) NE SR 1344, Nantahala National Forest; 16 1,

1 6 II, 3 j 6, 4 9, 1 j 9 (NCSM C-2175), 2 j 9, part of adult

exuvium (NCSM C-2178), 2 j 6,3] 9 (NCSM C-2180).

Cambarus parrishi was the only crayfish found at site (1), but

at site (2) nine C. b. bartonii also were collected. Of the four C.

parrishi taken at site (1), one of the females exhibited exoskeleton

decalcification and the male was very soft. At site (2), two males and

one female showed decalcification, two of the females were

soft, and part of an adult exuvium was found.

Cambarus (Puncticambarus) reburrus Prins

This species is endemic to headwater streams of the Savannah

and French Broad river basins, in Buncombe, Henderson, Jackson,

and Transylvania counties in the Blue Ridge province of North

Carolina (Cooper and Cooper 19776:214). There is some confusion,

however, about the precise distribution of the species. Its type locality

is a "Small tributary to the Horsepasture River from Sapphire

(= Fairfield) Lake off U. S. 64, 5.5 miles east of Cashiers, Jackson

County, North Carolina. Sapphire Lake is an impoundment from

which the tributary . . . flows several hundred yards into the Horse-

pasture River (upper Savannah River drainage)" (Prins 1968:459). It

should be noted that Fairfield Lake, although also on a tributary of

the Horsepasture River (Long Branch), is about 1.0 air mi (1.6 air

km) northwest of Sapphire Lake.

The type locality was the only known site for the species at

the time of its description, but four years later Prins and Hobbs (1972:

412) added the French Broad River basin to its range and suggested

that the "population described by Prins in the headwaters of the

Savannah River is an introduced one." Ross (1971:29-32), however,

had earlier discussed evidence for a probable Pleistocene breach of

the Blue Ridge by Savannah basin headwaters, and "an obvious encroach-

North Carolina Crayfishes 103

ment of Toxaway River of the Savannah River system upon the head-

waters of the French Broad River of the Tennessee River basin." Ross

further commented that "Toxaway Creek has not only breached the

Blue Ridge but it has also begun to drain some of the 2900 to 3000-

foot surface around Lake Toxaway, which once must have been part

of the French Broad basin." Later, Hobbs and Peters (1977:11) provided

evidence, based on the distributions of fishes and entocytherid

ostracods, that "the possibility of a former connection between parts

of the two basins exists." Despite these analyses, however, current

evidence indicates that Prins and Hobbs (1972:412) may have been

correct in suggesting that C. reburrus is not native to the Savannah

River basin.

The following additional collections of this species are all

from the same locality in the general area of the type locality.

Jackson Co. — pond in floodplain Horsepasture R along US 64, 1.2

rd mi (1.9 rd km) W jet SR 1119; 1 9 with young (NCSM C-890),

1 ovig 2 (NCSM C-891), 5 Jun 1977, ALB, 3 $ with young

(NCSM C-253, 258, 259), 1 j $ , 1 S II (NCSM C-260), 4 6 II, 1 $

1 j 9 (NCSM C-912), 27 Jun 1977, DLS, ALB. Several collections

made elsewhere in the Horsepasture watershed yielded only C. b.

bartonii.

At all localities where C. reburrus has been taken, it has proved

to be a creature of slack or slow-moving waters of low gradient and

generally with considerable organic debris. To our knowledge, it has

not been taken in swift, high-gradient streams devoid of detritus.

Collections we made in September 1984 in two such headwater streams

of the Chatooga River south of Highlands, Macon County, streams

that in North Carolina are independent of the more eastern head-

waters of the Savannah, produced only C. asperimanus. Suitable C.

reburrus habitat is unlikely in the lower Horsepasture and Toxaway

rivers, and in the Thompson River. It almost certainly exists, however,

in those parts of the Horsepasture watershed between Sapphire and

Fairfield lakes and the northern limits of Cashiers, in the headwaters

of the Chatooga River in and around Cashiers, in the floodplain of the

Toxaway River around Lake Toxaway above Toxaway Falls, and in

the Whitewater watershed above Whitewater Falls. We can find no

evidence that C. reburrus has been taken from any of these areas, but

perhaps more field work will reveal its presence in one or more of

them.

Although fairly common and widespread in the upper reaches

of the French Broad River basin, C. reburrus has yet to be verified

any farther downriver in this system than eastcentral Buncombe

104 J. E. Cooper and A. L. Braswell

County (see Prins and Hobbs 1972:412 for localities near Black

Mountain and Swannanoa, and the additional Buncombe County

locality listed below). Prins and Hobbs (1972:412) reported the species

from "Madison County — 3.5 mi. west of Old Fort in Pisgah National

Forest on U. S. Hwy. 70, . . . 17 June 1940, J. C. Moore, coll.," and

added, "These specimens are somewhat atypical and are thus

tentatively assigned to this species." Hobbs (1989:27) included

Madison County in the range of the species. The locality in

question, however, is in McDowell County and in the headwaters of

the Catawba River basin, not the French Broad. Old Fort is over 5 air

mi (8 air km) east of the Buncombe County line and the nearest

headwater creeks of the Swannanoa River (French Broad). The

Madison County line is some 24 air mi (38 air km) west of Old Fort,

so Madison County is in error. Hobbs very kindly loaned us the

specimens collected over 50 years ago at this site (USNM 131904; 1

j 8, 3 j $). Comparison with C. reburrus of similar sizes and sexes

showed that the questionable specimens did not belong to this

species. Hobbs later pointed out (in litt.) that they may belong to an

undescribed Puncticambarus that occurs in the Catawba River

basin, one of those subsumed under C. (P.) sp. C.

Further doubt is cast on the occurrence of C. reburrus in lower

parts of the French Broad basin by collections we made in July and

September 1984. We collected over 300 specimens at 11 sites of

various kinds in the Pisgah National Forest in Madison County, and

78 specimens at 4 sites in the Pigeon River drainage in Haywood

County, but found no C. reburrus. Our current knowledge, then,

indicates that this species is absent from the Pigeon (and probably

Nolichucky) hydrologic units of the French Broad, and from the main

French Broad and its tributaries northwest of the Asheville area.

Hobbs and Peters (1977:36) reported the ostracod, Dactylocy there

megadactylus Hart and Hart, an ectocommensal of several crayfish

species, from 37 localities in North Carolina, "in which it is confined

to the Piedmont Plateau." They inadvertently listed C. reburrus among

the ostracod's hosts, but their distribution map for D. megadactylus

(Hobbs and Peters 1977:71) showed no localities within the range of

this Blue Ridge crayfish.

So many additional collections of C. reburrus have been

made from Henderson County, particularly from the French Broad

and Mills rivers, that we see no reason to list them. The following

localities, however, are of sufficient interest to be specified: Buncombe

Co.— (1) Swannanoa R at SR 2416, 0.1 rd mi (0.2 rd km) S jet

SR 2002, ca. 1.5 air mi (2.4 air km) NW Wilson; numbers and sexes

North Carolina Crayfishes 105

not available (NCSM C-2194), 25 Sep 1984, ALB, JEC. Transylvania

Co.— (2) French Broad R at River Mile (RM) 195.8 along Wilson Rd

at head Elm Bend; no other data available, M. Ford; (3) French

Broad R at mouth Williamston Crk (RM 192.8) at US 276; 1 9

(NCSM C-1686), 23 Jun 1977, MF; (4) West Fk French Broad R near

jet SR 1309 & 1312, ca. 4.2 air mi (6.7 air km) NW Rosman; 1 9

(NCSM C-2221), 18 Oct 1984, V. Schneider; (5) Davidson R at US

276, 0.2 rd mi (0.3 rd km) SE of FSR to Pisgah Forest National Fish

Hatchery, ca. 4.3 air mi (6.9 air km) NNW cntr Brevard; 1 9, 13 Aug

1993, J. Weems.

Orconectes {Crockerinus) erichsonianus (Faxon)

Hobbs (1981:263) listed this species and C. latimanus as

associates of C. hiwasseensis at a locality in the Hiwassee River

system of North Carolina. This is the only published record for O.

erichsonianus in North Carolina. Hobbs (in litt.) said, "the locality is

1.6 miles west of the junction of U. S. Hwy. 64 on State Route 60,

Cherokee County — Nottely River, collected on June 12, 1960, by K.

W. Simonds . . . ." He added, "the whereabouts of the specimens is

a puzzle. They were never catalogued and perhaps are still shelved

with the uncatalogued material here [at the U. S. National Museum

of Natural History]."

On 21 July 1984, we collected 47 crayfish at a site that must

have been the same as, or very close to, the one in question — the

Nottely River at NC 60, 2 air mi (3.2 air km) SSW of Ranger.

This collection contained only C. b. bartonii, C. hiwasseensis, and

an unidentified Cambarus. Collections that we made on several

occasions elsewhere in the Hiwassee River basin in Cherokee and

Clay counties also failed to yield any Orconectes, and the species is

not known from this system in Georgia (Hobbs 1981:21, 287). Hobbs

later (1989:35, 84, 89) did not include North Carolina in the general

range of O. erichsonianus, and we are convinced that the species does

not occur here. It could have been extirpated from the State, but its

historic range just west of North Carolina is limited to the Appalachian

Plateau and the Ridge and Valley physiographic provinces, no

elements of which are found in North Carolina.

Orconectes {Crockerinus) virginiensis Hobbs

This is one of only two species of Orconectes known to occur

on the Atlantic versant in North Carolina, where it reaches the southern

limits of its range. It is also the only member of subgenus Crockerinus

106 J. E. Cooper and A. L. Braswell

in the State. Its type locality is a tributary of the Nottoway River,

which is a major trunk of the Meherrin River, in southeastern Virginia

(Hobbs 1951:124-125). The Meherrin River joins the Chowan River

at the Gates County, North Carolina, line. Orconectes virginiensis

originally was known in the Chowan basin of North Carolina from

five specimens collected at a single locality in Hertford County:

Cutawhiskie Creek (Swamp), a tributary of Potecasi Creek, 1 air

mi (1.6 air km) southwest of Menola (Cooper and Cooper 1977c:

215). The collection, made on 13 August 1974 by Chris Ellis, consisted

of a form I and two form II males (NCSM C-33), and two females

(NCSM C-31, 32). A sixth Chowan basin specimen, an adult female,

was collected on 24 July 1985 from a submerged log in the Meherrin

River at CR 1175 (Parkers Ferry), downstream from the confluence of

Potecasi Creek, about 3.2 air mi (5.1 air km) north-northwest of the

center of Winton, Hertford County, by David R. Lenat.

Although Hobbs (1989:38) gave the general range of O.

virginiensis as "Chowan drainage system in North Carolina and

Virginia," Cooper and Cooper (1977c:215) had reported it earlier from

the lower Roanoke River basin in Martin County. A form I male

(USNM 116979) was collected in Ready Branch, a tributary of

Sweetwater Creek, 4.5 air mi (7.2 air km) south of Williamston, on

28 March 1949 by E. C. Raney. This species is now known from the

following additional localities in the Roanoke River basin: Bertie

Co.—(l) Roanoke R at NC 45/308 at Washington Co line, 5.6 air mi

(9.0 air km) ENE cntr Sans Souci; 1 9, 2 j 9, 6 Jul 1986, DRL.

Halifax Co.— (2) Roanoke R at NC 258 at Northampton Co line,

2.9 air mi (4.6 air km) E cntr Spring Hill; 1 j 6, 9 Jul 1987, DRL.

Martin Co. — (3) Ready Br at US 17 bridge, ca. 5.5 air mi (8.8 air

km) S Williamston; 1 j 9 (NCSM C-363), 17 Jun 1980, JP.

The adult female collected at the Bertie County site was found

in a submerged log. About this locality, which is not far west of

Batchelor Bay, a brackish estuary of Albemarle Sound, Lenat (in litt.)

said that the water there was about 200 m wide, slow-moving, and

"may be slightly brackish." He added, "The shore areas have abundant

growths of water lily, with many pieces of dead wood on the bottom.

Prolific algal growths suggested some enrichment this year. The fauna

is primarily freshwater, but also contains some estuarine taxa." At the

Martin County site, a form I male and two female P. a. acutus were

collected with the O. virginiensis.

Occurrence of this species in both the Chowan and lower

Roanoke basins is not surprising, since during the Pleistocene the

Chowan "would have been a tributary of the Greater Roanoke

North Carolina Crayfishes 107

River" (Lachner and Jenkins 1971:62). Little has been reported on the

distribution and natural history of this species, especially in

North Carolina, and it remains one of the State's least known and

ostensibly rarest crayfishes.

Orconectes (Procericambarus) spinosus (Bundy)

There are no published localities for the occurrence of any

member of subgenus Procericambarus, and specifically for O. (P.)

spinosus, in North Carolina. Hobbs (1981:297) indicated its general

range as "Streams in the Coosa and Tennessee river basins in

Alabama, Georgia, North Carolina, Tennessee, and Virginia," but later

(Hobbs 1989:50, 85, 89) did not include North Carolina within the

range of the species. Prior references to Cambarus (= Orconectes)

spinosus in the Tar River (Faxon 1890:632; Harris 1903: 130, 142;

Ortmann 1905:115, 1931:87, 88), all apparently based on specimens

reported by Bundy (1877) from Rocky Mount, refer to the undescribed

Orconectes sp. A.

North Carolina specimens assigned to O. spinosus have been

collected at the following localities in the Cheoah River watershed

of the Little Tennessee River basin: Graham Co. — (1) Tulula

(Talula, Talulah) Crk at SR 1211, ca. 0.3 air mi (0.5 air km)

E Robbinsville; 1 6 II, 1 9- (NCSM C-252), 25 Nov 1978, DLS,

ALB; (2) Cheoah R just upstream from sewage treatment plant

outfall Robbinsville; 5 6 II, 3 9 (NCSM C-389), 8 Oct 1980, J. H.

Davies; (3) Tulula Crk at NC 143, 0.3 air mi (0.5 air km) E cntr

Robbinsville; 2 6 I, 2 6 II, 4 9 (NCSM C-1213), 23 Oct 1982,

RWV, ALB, D. Sever, 5 ovig 9 (NCSM C-2302), 2 8 I, 1 6 II, 19

j 6, 9 j 9 (NCSM C-2303), 25 Apr 1985, ALB; (4) Tulula Crk at

SR 1138, just N Robbinsville; 7 6 I, 1 6 II, 5 j S, 3 j 9 (NCSM

C-2306), 2 ovig 9 (NCSM C-2312), 2 ovig 9 (NCSM C-2313), 4 6

I (NCSM C-2352), 26 Apr 1985, ALB.

At locality (2), three of the form II males showed considerable

exoskeleton decalcification, and one of them and another form II

male and one female were soft. The specimens taken at locality (3)

were found under rocks in the creek, and the species appeared to be

more abundant than the only other crayfish found there, C. b. bartonii.

The two species also occurred together at locality (4).

Orconectes (Procericambarus) sp. B

A large species of Orconectes, previously considered to be

introduced Orconectes juvenilis (Hagen) [= Orconectes (Proceri-

108 J. E. Cooper and A. L. Braswell

cambarus) rusticus (Girard)] by Hobbs and Walton (1966) and

Hobbs et al. (1967), occurs in the New-Kanawha basin of Virginia

(Hobbs, in litt.). This animal also has been taken in New River

headwaters at the following North Carolina localities: Alleghany Co. —

(1) small stream entering New R just E US 21-221, ca. 0.2 air mi

(0.3 air km) S Virginia state line, 3.0 air mi (4.8 air km) NNE jet US

21 & 221 at Twin Oaks; 1 ovig 9 (NCSM C-284), 20 May 1978,

REA, Jr, JEC. Ashe Co.— {2) South Fk New R at SR 1602, ca. 0.6 air

mi (1.0 air km) SE jet US 221, 2.6 air mi (4.2 air km) ENE town

Nathans Creek & 7.8 air mi (12.4 air km) NE cntr Jefferson; 3 8 II, 2

j 8, 4 9 (NCSM C-257), 20 Jul 1978, DSL; (3) South Fk New R at

E side NC 88 bridge and along SR 1588, ca. 0.5 rd mi (0.8 rd km)

NE Orion & 4.8 air mi (7.6 air km) E West Jefferson; 2 8 I, 1 8 II

(NCSM C-276), 1 ovig 9 (NCSM C-277), 1 8 II (NCSM C-256),

21 May 1978, REA, Jr, JEC; (4) South Fk New R along CR 1566,

0.3 rd mi (0.5 rd km) S jet SR 1567; 2 ovig 9 (NCSM C-803, 804),

4 8 II (NCSM C-805), 20 May 1978, REA, Jr, JEC; (5) South Fk

New R at NC 221, ca. 1.7 air mi (2.7 air km) WSW Scottsville; 1 j 8,

2 9 (NCSM C-2358), 27 May 1985, F. Winborne, DRL; (6)

Buffalo Crk near confluence Little Buffalo Crk, ca. 2.5 mi (4.0 km)

W cntr West Jefferson; 1 j 8 (NCSM C-2355), 30 May 1985,

DRL, FW; (7) Dog Crk at SR 1592, 4.0 mi (6.4 km) E Jefferson; 1 j

9 (NCSM C-508), 6 Aug 1978, J. R. Clamp, M. Dennis.

This species was very common at locality (3), where it occurred

in close association with another large crayfish, Cambarus {Hiati-

cambarus) chasmodactylus James. The two often were found under

the same cover, usually medium-size to large rocks in cold, fast-

moving water, and in the same gravel riffles. Both species also

occurred together at locality (2). At locality (4), the Orconectes was

collected with C. chasmodactylus and C. robustus.

Procambarus (Ortmannicus) medialis Hobbs

Procambarus {Ortmannicus) pearsei (Creaser)

Procambarus {Ortmannicus) plumimanus Hobbs and Walton

These three species comprise a "disjunct" enclave at the northern

periphery of the range of the Planirostris Group of the genus

Procambarus, a fact that is of considerable importance in under-

standing the evolutionary history of the group. Hobbs (1975:15)

summarized their distributions: "The Neuse River basin appears to

mark the southern limit of the range of P. medialis, the northern

[Northeast] Cape Fear River basin marks the southwestern limit of

the range of P. plumimanus, and the range of P. pearsei encom-

North Carolina Crayfishes 109

passes most of the coastal plain lying in the Cape Fear River basin

(excluding the Northern [Northeast] Cape Fear) southward through

the Little Pee Dee River basin." The following records and discussions

support this statement, slightly expand and clarify the known ranges

of these species, add some information on their natural history, and

help explain a distributional anomaly for P. plumimanus.

Procambarus medialis

As mentioned by Cooper and Ashton (1985:10), although the

type locality of P. medialis is in the Tar-Pamlico basin in Halifax

County (Hobbs 1975:13), this and one other site near it in the drainage

of Deep Creek are the only places within this basin where the species

currently is known to occur. All other collection sites are within the

Neuse River basin. Eleven specimens were captured in wire minnow

traps at three localities in the Neuse basin during the N. lewisi study

(Braswell and Ashton 1985), and one of them was in the mainstem

Neuse River in Lenoir County. Identical traps were employed at over

180 stations in the Tar-Pamlico basin, from its headwaters in Person

County to central and southern Beaufort County, but yielded no

additional P. medialis. One of these stations was in Beech Swamp,

less than 5 air mi (8 air km) west of Deep Creek, and another was

in the Tar River near its confluence with Deep Creek in Edgecombe

County. There are no obvious ecological, edaphic, or physiographic

reasons, however, for this apparent severe restriction of the known

range of P. medialis in the Tar-Pamlico basin. During the N. lewisi

survey, little collecting was done in roadside ditches or isolated lentic

waters, and no digging of burrows was attempted. More thorough

sampling of these possibly preferred habitats might reveal a wider

distribution pattern for P. medialis in this river basin, although it

also inhabits larger, flowing waters in the Neuse basin.

The following additional localities are in the Neuse River basin:

Johnston Co.— {!) Black Crk at NC 50, 5.5 air mi (8.8 air km) NNE

Benson; 1 9 (NCSM C-144), 27 Jan 1979, trap, K. Everett; 1 9

(NCSM C-230), 30 Jan 1979, trap, KE; 1 9 (NCSM C-140), 10 Feb

1979, trap, KE; (2) Hannah Crk at US 701, 10 air mi (16 air km)

E Benson; 4 9 (NCSM C-1079), 22 Apr 1979, trap, P. S. Freed, 1 6

II, 1 9, 1 j 9 (NCSM C-1097), 23 Apr 1979, trap, PSF; (3) dry

roadside ditch, Truck Lane and East Rose St, Smithfield; 1 c? I, 1

6 II, 1 9 (NCSM C-2270), 17 Apr 1985, W. M. Palmer, ALB.

Lenoir Co.— (4) Neuse R at NC 55, 1.1 rd mi (1.8 rd km) E jet SR

1809, 5.2 air mi (8.3 air km) NE Kinston; 1 6 II (NCSM C-231), 29

Jan 1979, trap, PSF.

110 J. E. Cooper and A. L. Braswell

At locality (3), each of the three specimens was found under

a separate railroad crosstie along the dry ditch, near entrances to

burrows that could be seen to reach water. The ditch normally

contains water, but this area was in the midst of a prolonged drought

that had been broken only by some rainfall on the previous night.

Other ditches had some water. In captivity, the form I male, collected

in April, molted and died on 29 September 1985, but remained

form I.

Procambarus pearsei

Hobbs (1975:14) mapped 22 unspecified localities for P. pearsei

in the Cape Fear, Lumber, and Waccamaw basins in Bladen, Brunswick,

Columbus, Cumberland, Robeson, and Sampson counties. This same

map inadvertently indicated an additional locality for P. pearsei in

the Neuse River drainage of Johnston County. This should have been

for P. medialis; the position of the dot corresponds to a locality for

this species given by Hobbs (1975:13), ". . . 5.3 miles south of

Smithfield on U. S. Highway 706." The nearest headwater stream of

the Cape Fear basin, Mingo Swamp, rises at the extreme western edge

of Johnston County, about 2.5 air mi (4.0 air km) SSW of site (1)

above. Thus, the range of P. pearsei given by Hobbs (1989:70) as

"Johnston and Sampson counties, North Carolina, south to . . . .,"

currently requires the deletion of Johnston County.

The following localities add P. pearsei to the Cape Fear basin in

Bladen County, and extend the known range slightly west in the

Lumber River basin into Hoke and Scotland counties. CAPE FEAR

RIVER BASIN. Bladen Co.— (I) pond along SR 1327, 1.6 rd mi (2.6

rd km) NW jet SR 1325, 5.7 air mi (9.1 air km) W Ammon, in

vicinity Little Singletary and Horseshoe (Suggs Mill Pond) lakes; 1 6

I, 1 j 6 (NCSM C-304), 17 Mar 1979, P. S. Ashton, REA, Jr., ALB;

(2) Horseshoe Lake, 0.7 rd mi (1.1 rd km) NE SR 1327; 1 6 I, 8 j 6,

1 9, 6 j 9 (NCSM C-1602), 23 Feb 1980, PSA, REA, Jr, NCSM

Jr Curators. LUMBER RIVER BASIN. Hoke Co.— (3) Antioch Bay

off NC 211, 1.7 air mi (2.7 air km) SSE Antioch; 4 8 I, 3 9, parts of

9 (NCSM C-2240), 1 Feb 1985, RWL, DLS, DFL; (4) pools in

powerline cut Antioch Church Bay, E side NC 211, 0.6 rd mi (1.0 rd

km) SSE jet SR 1447, 2 air mi (3.2 air km) SSE Antioch; 1 6 II, 2

9 (NCSM C-2241), 23 Feb 1985, DLS, RWL, DFL. Scotland Co.—

(5) borrow pit near jet SR 1400 & 1413, ca. 3.5 air mi (5.6 air km)

NW Wagram; 1 S I, 1 j 6\ 2 j 9 (NCSM C-2011), 8 Oct 1974,

MRC, WMP, ALB, JEC.

North Carolina Crayfishes 111

The remains of a form I male and at least one other P. pearsei

(NCSM C-1277) were removed from the stomach of a redtailed hawk,

Buteo jamaicensis, found dead near Hallsboro, Columbus County, on

8 April 1983, and prepared as a museum specimen by Gilbert Grant.

Seven females carrying late-instar young (NCSM C-2242-2244, 2246-

2249) were found at a site in Robeson County on 23 February 1985

by DLS, RWL, and DFL. A female (NCSM C-1951) was found alive

on NC 211, about 3 air mi (4.8 air km) south of Bolton, Columbus

County, at night on 24 March 1975.

Procambarus plumimanus

Hobbs and Walton (1958:10-11) provided a locality for P.

plumimanus in Craven County and two in Duplin County, and Hobbs

(1975:14) mapped two localities in Duplin County, three in Craven

County, one in Carteret County, and one apparently on the Craven-

Carteret line. Hobbs (1989:70) added Jones County to the range of

the species, but gave no specifics since this was a general checklist.

The following localities add to our knowledge of the distribution

of this crayfish. NORTHEAST CAPE FEAR RIVER BASIN. Pender

Co. — (1) borrow pit "Back Island," Holly Shelter Game Lands, 0.3 mi

(0.5 km) N Lodge Rd; 1 6 II, 4 j 6,1 9, 1 j 9 (NCSM C-870), 4 j

8,5 j 9 (NCSM C-871), 3 Oct 1981, REA, Jr; (2) borrow pit Lodge

Rd, Holly Shelter Game Lands, ca. 2 air mi (3.2 air km) NW US 17,

5.8 air mi (9.3 air km) SW Edgecombe; 3 6 I, 2 j 6\ 1 9 (NCSM C-

872), 24 Jul 1981, REA, Jr; (3) dr of Harrisons Crk off NC 210, ca. 1

rd mi (1.6 rd km) N jet SR 1574, 4.2 air mi (6.7 air km) NW

Hampstead; 1 6 I, 1 j 8,2 9 (NCSM C-933), 13 Mar 1977, E.

Flowers, DLS. NEW (WHITE OAK) RIVER BASIN. Carteret Co.—

(4) borrow pit along SR 1125, 5.3 mi (8.5 km) NW Newport

(vicinity Northwest Prong Newport R); 1 9 (NCSM C-1942), 25

Mar 1975, MRC, JEC; (5) slough along Millis Rd (SR 1112

extended), 0.7 mi (1.1 km) W jet SR 1124, 5.5 air mi (8.8 air km)

WSW Newport (Jason Br of Southwest Prong Newport R); 2 6 \,

1 6 II, 1 j 6,1 9, 1 j 9 (NCSM C-2097), 25 Mar 1975, MRC,

ALB, JEC. Jones Co.— (6) drainage ditch along FSR 152 (Black

Swamp Rd), ca. 7 mi (11.2 km) ESE Maysville, 1 8 I (NCSM C-

294), 7 Apr 1979, DSL; (7) slough on Great Lake Rd (SR 1101

extended as FSR 126), ca. 10 air mi (16 air km) SE Maysville

(Hunter Crk dr); 2 6 I, 1 6 II, 1 j 6, 1 j 9 (NCSM C-2069), 26 Mar

1975, MRC, JEC. New Hanover Co. — (8) cypress pond at Carolina

Beach State Park; 1 6 I, 1 9 (NCSM C-850), 3 May 1980, PSA,

112 J. E. Cooper and A. L. Braswell

REA, Jr; this locality is on a narrow peninsula between the Cape Fear

River and Onslow Bay. Onslow Co. — (9) permanent ditch and pools

along US 17, 0.4 mi (0.6 km) N jet SR 1103, ca. 3.7 air mi (5.9 air

km) S Verona; 1 6 I, 1 6 II, 4 $ (NCSM C-70), 10 Aug 1976, M.

M. Browne.

Although the natural range of P. plumimanus appears to be

limited to the Northeast Cape Fear and New (White Oak) basins, and

this species did not appear at any of the N. lewisi survey stations

sampled in the Neuse basin (Cooper and Ashton 1985:10), two

localities for the species within the lower Neuse basin are known.

The type locality, "Roadside ditch 2.2 miles southeast of Havelock,

Craven County, North Carolina on Hwy. 70" (Hobbs and Walton 1958:

10) apparently lies close to East Prong Slocum Creek and not far

from the head of Hancock Creek, both north-flowing streams that

drain into the Neuse River on either side of the Cherry Point Naval

Reservation. Only swamps separate this area from nearby headwaters

of the Newport River not far to the south. The second locality, also in

the Neuse basin in Craven County, is a ditch along Catfish Lake Road

(SR 1100), 3.0 air mi (4.8 air km) southwest of the town of Croatan,

in the drainage of East Prong Brice Creek.

This apparent expansion of the range of P. plumimanus into the

lower Neuse basin possibly can be explained by the fact that in

Carteret, southern and eastern Craven, and southern Jones counties,

as in much of the poorly drained outer (tidewater) Coastal Plain,

contemporary drainage distinctions have become blurred. The drainage

divides, very low to begin with, have been breached by stream

channelization, a vast system of man-made drainage canals, and the

Intracoastal Waterway. Drainage canals along SR 1100 east and south-

east of Catfish Lake provide access from the White Oak River basin

to West Prong Brice Creek of the Neuse basin. Farther east, the

Harlowe (Clubfoot), Adams Creek, and other canals, link elements of

the Neuse River with the Newport and North rivers. Precisely what

roles the easternmost of these interbasin waterways might play in

crayfish distributions will not become clear until we have an under-

standing of the tolerance that various species display for the physico-

chemical features of their waters. Some crayfishes, notably P. a. acutus

and Fallicambarus (Creaserinus) fodiens (Cottle), and even C. diogenes,

are widely distributed in tidewater North Carolina. They often occur

in close proximity to saline estuaries and tidal creeks, and there is a

population of F. fodiens across Croatan Sound on Roanoke Island.

Interbasin connectors also are present in other coastal river basins

(see below).

North Carolina Crayfishes 113

THE HYDROLOGIC UNITS AND THEIR CRAYFISH FAUNAS

Seventeen major river basins generally are recognized within

North Carolina (Heath et al. 1975:152). All of them include smaller

hydrologic units, some of which are autonomous drainage systems

within the State. All but five of the major river basins lie within the

huge drainage of the Atlantic Ocean, and flow generally east and

south to empty into broad, saline estuaries and sounds of the ocean.

The exceptions are (1) the Hiwassee, Little Tennessee, French Broad,

and Watauga rivers, which drain north and west into the Tennessee

River, and (2) the New River, which drains north and west via the

Kanawha River into the Ohio River. These western North Carolina

waters ultimately flow into the Mississippi River drainage of the Gulf

of Mexico. In North Carolina, the west-flowing and east-flowing

montane headwaters of river systems are separated by the Appalach-

ian (Eastern Continental) Drainage Divide, represented in the State

by the Blue Ridge physiographic province.

The Waccamaw River and its tributaries, along with Lake

Waccamaw, are recognized as a hydrologically and faunistically

distinctive drainage unit by some biologists (see Bailey 1977:

269, 273; Shute et al. 1981:18-22), and are so treated here. Another

hydrologic unit, the Northeast Cape Fear River, probably also should

be considered an autonomous drainage system, because it does not

join the Cape Fear River until it reaches the estuary at Wilmington.

In addition, the Northeast Cape Fear is faunistically distinct, apparently

lacking over a dozen species of lowland freshwater fishes found

in the Coastal Plain portions of the Cape Fear itself (Rohde et

al. 1979:114-115: Menhinick 1991). The Northeast Cape Fear also

lacks two crayfish species found in the lower Cape Fear, but has one

species that is absent from the Cape Fear (see below). Historically as

well as faunistically, the Northeast Cape Fear appears to have more

in common with the New (White Oak) drainage system than with the

Cape Fear. During the Pleistocene, when sea level attained its

highest stand (as indicated by the Surry Scarp, which marks the marine

terrace formed by the Wicomico Sea), much of the southeastern

Coastal Plain of North Carolina was covered by the ocean (Rohde

et al. 1979:113, 116). This inundated area included today's New

(White Oak) River drainage, most of the Northeast Cape Fear, and

part of the lower Cape Fear. The early coastal Cape Fear and North-

east Cape Fear basins were at that time broadly separated, and they

are still separate systems today.

114 J. E. Cooper and A. L. Braswell

Tennessee-Ohio-Mississippi

Hiwassee River — The Hiwassee River system rises in the Blue

Ridge of northcentral Georgia, and in Clay and Cherokee counties,

North Carolina. It flows generally north and west across the southwest

corner of the State and into Tennessee, where it merges with the

Tennessee River. Two streams of this system head in the southwest

corner of Cherokee County and flow south into the Toccoa River in

Georgia, part of the Ocoee River system. The Ocoee then joins the

Hiwassee in Tennessee.

Cambarus hiwasseensis and C. parrishi are upper Hiwassee

endemics, occurring in both Georgia and North Carolina. The type

locality of C. hiwasseensis is a tributary to Peachtree Creek, 0.8 mi

(1.3 km) north of Peachtree School on US 64A, Cherokee County,

North Carolina (Hobbs 1981:260-261). Other species that occur in

the Hiwassee system in North Carolina but have broader ranges are C.

b. bartonii, C. nodosus, and C. acanthura. The latter two species

are known in North Carolina only from the Hiwassee. Hobbs (1981:

20, 22) reported C. latimanus in this system in Georgia, and said

(Hobbs 1981:263) that it was found with C. hiwasseensis and O.

erichsonianus at a locality on the Nottely River in North Carolina. In

addition, his range map for C. latimanus (Hobbs 1981:115) showed

Georgia localities west of Nottely Lake in Union County, and in

Fannin County just west of the North Carolina state line. Further-

more, Bouchard (1972:36) recorded the species "downstream in the

Ocoee (Tennessee) River drainage to the Little Frog Mountains in

Polk County, Tennessee." This area is in the Hiwassee basin just west

(downriver) of North Carolina. Thus, C. latimanus, which in

North Carolina is a species of the eastern Piedmont Plateau and the

Coastal Plain, also may occur in the Hiwassee basin of the Blue

Ridge in at least Cherokee County. It did not turn up, however, in

our Hiwassee field work, which included the Nottely River. Hobbs

and Peters (1977:27) cited a record for Cambarus (Hiaticambarus)

longirostris Faxon in the Hiwassee basin in Cherokee County, "Valley

River at Andrews . . . (Crawford 1961:244)," and Hobbs (1981:171-

172) found this species in the headwaters of the Nottely River in

Union County, Georgia, which is a considerable distance upriver from

North Carolina. The distribution maps in James (1966:12-13), however,

showed no localities for this species within the Hiwassee basin in

North Carolina. In July and September 1984, we failed to collect

this crayfish at any site within the system, including the Valley

River near Andrews. Nevertheless, C. longirostris could occur in the

Hiwassee basin in North Carolina. Cambarus carolinus also may occur

North Carolina Crayfishes 115

there, but specific records are lacking. There is a small, forest-green

Cambarus in Cherokee County that is as yet unidentified.

Little Tennessee River — The main artery of the Little Tennessee

River originates in the Blue Ridge of Rabun County, Georgia. Its

tributaries drain most of Macon County, and parts of Swain and

Graham counties, North Carolina. Its major eastern tributary, the

Tuckasegee River, heads in Jackson County and flows northwest into

Fontana Lake. Another eastern tributary, the Cullasaja River, begins

in southeastern Macon County and joins the Little Tennessee River

south of Franklin. A western tributary, the Cheoah River, heads in

southern Graham County and flows northwest through Santeetlah

Lake, to merge with the Little Tennessee River at Cheoah Dam near

the state line. A second major western tributary, the Nantahala

River, rises in western Macon County and along the eastern border of

Clay County, and flows primarily north into Fontana Lake. Beyond

this lake the Little Tennessee flows west into Tennessee, joining the

Tennessee River near Lenoir City.

One crayfish, C. georgiae, is endemic to the Little Tennessee,

where currently it is known from two sites along the main river in

Rabun County, Georgia, and several lower elevation sites in the Cullasaja

River watershed of North Carolina. More wide-ranging species of this

system are C. b. bartonii, C. longirostris, and C. asperimanus. Specific

localities for C. carolinus are reported here, and Hobbs and Peters

(1977:56) referred to a C. (J.) sp. in Jackson County (Tuckasegee

watershed), which may turn out to be C. carolinus or an undescribed

species. Orconectes spinosus is herein reported from the Cheoah

River watershed of this basin.

French Broad River — The uppermost tributaries of the French

Broad River rise on the western slopes of the Blue Ridge in Transylvania,

Henderson, and Buncombe counties. Its major western tributary, the

Pigeon River, heads in western Haywood County and is an autonomous

unit in North Carolina. However, it joins the French Broad near Newport,

Cocke County, Tennessee. The main eastern tributaries, the Cane,

Nolichucky, and Toe rivers, rise in Avery, Mitchell, and Yancey counties.

They, too, form an independent unit that flows northwest and enters

Tennessee as the Nolichucky. This river then turns west and merges

with the French Broad at the upper end of Douglas Lake. The French

Broad River initially flows northeast in North Carolina, then turns

northwest and enters Tennessee about 5.5 air mi (8.8 air km) downriver

from Hot Springs, Madison County. It joins the Holston River at

Knoxville to form the Tennessee River.

116 J. E. Cooper and A. L. Braswell

The endemic North Carolina crayfish, C. reburrus, is distributed

throughout the upper French Broad, but appears to be absent from the

Pigeon (and probably Nolichucky) watersheds, and from the main

French Broad system northwest of Asheville. The other species of

this river basin are C. b. bartonii, C. longirostris, C. asperimanus, C.

dubius, C. robustus (Hobbs and Peters 1977:30), and C. sp. C. The

type locality of C. asperimanus is Flat Creek, Montreat, Buncombe

County (Faxon 1914:391). We collected many specimens of what may

be an undescribed Puncticambarus in three streams in Madison County,

but for now these are assigned to C. sp. C. Bouchard (1978:36) reported

a specimen of C. latimanus from the French Broad in North Carolina,

extralimital to any part of the known range of the species, but considered

it probably an introduction.

Watauga River — The Watauga River rises in western Watauga

County west of Grandfather Mountain and flows north and west into

Tennessee. A southwestern tributary, the Elk River, heads in

northern Avery County. It is a minor independent subdrainage in

North Carolina, but joins the Watauga in Tennessee. The Watauga is

impounded just beyond North Carolina's borders,' but eventually

merges with the Holston River of the Tennessee River system.

The crayfish fauna of the Watauga River basin in North Carolina

is not well known, but consists of C. b. bartonii, C. longirostris, C.

asperimanus, C. dubius, and C. robustus.

New River — The two major western tributaries of New River —

North Fork New River and South Fork New River — head in Ashe and

Watauga counties, respectively. They flow northeast and merge at the

northern Ashe-Alleghany county line to form the New River just before

it enters Virginia. Eastern tributaries, Little River and Brush Creek,

rise in central and eastern Alleghany County, and flow northward

into Virginia to join the New River. The New River merges with the

Kanawha River near Charleston, West Virginia.

Cambarus chasmodactylus and probably Orconectes sp. B are

endemic to the New-Kanawha system, and in North Carolina they are

limited to, and close associates in, fast-flowing tributaries of the

New River. Records for these common species in the eastern tributaries

in Alleghany County, however, are sparse. The other members of the

New River crayfish fauna in North Carolina are C. b. bartonii, C.

asperimanus (Watauga County), C. dubius, and C. robustus.

Atlantic

Savannah River — The Savannah River begins as fast-flowing

streams in deep ravines on the eastern slopes of the Blue Ridge

in southeastern Macon, southern Jackson, and southwestern

North Carolina Crayfishes 117

Transylania counties. In North Carolina, its easternmost tributaries —

the Toxaway, Horsepasture, Thompson, and Whitewater rivers — flow

south as independent streams, entering Lake Jocassee and the Keowee

River system of the Savannah basin not far into South Carolina. West

of these rivers, the Chatooga River and several even more western

tributaries of that river head in Macon County and flow independently

south. The Chatooga River forms part of the upper boundary between

Rabun County, Georgia, and Oconee County, South Carolina. The

Tallulah River, which rises in the southeastern corner of Clay County

on the slopes of Standing Indian Mountin, flows south into Towns

and Rabun counties, Georgia, where it joins the Tugaloo River of

the upper Savannah system.

The crayfish fauna of the Savannah River basin in North Carolina

consists of the endemic C. reburrus (currently known in this system

only from the floodplain of the Horsepasture River in Jackson County),

C. b. bartonii, C. asperimanus, and Cambarus sp. (Hobbs and Peters

1977:38). The type locality of C. reburrus is in the Savannah basin

(see the preceding account for this species). Little is known of

the crayfish fauna of the Tallulah River in North Carolina, but

C. asperimanus and C. b. bartonii undoubtedly occur there.

Broad River — This system rises in the west as headwater streams

in southern and eastern Henderson, western Polk, and eastern

Buncombe counties, and in the north in southern McDowell and

northern Rutherford and Cleveland counties. A western tributary, the

North Pacolet River, heads in southern Polk and Henderson counties,

and flows independently into the Pacolet River of South Carolina.

The Pacolet and Broad rivers merge at the juncture of Cherokee,

Union, and Chester counties, South Carolina. Eastern tributaries

head in eastern Cleveland and western Lincoln and Gaston counties,

flow independently into South Carolina, then soon join the Broad

River at the line between Cherokee and York counties. The Broad

flows through the eastern foothills and western Piedmont of North

Carolina, across South Carolina, and merges with the Saluda River at

Columbia to form the Congaree River. The Congaree joins the Wateree

River (continuation of the Catawba River), and the Wateree flows

into the Santee system via Lake Marion.

The crayfish fauna of the Broad River basin in North Carolina

consists of C. b. bartonii (the eastern and southern limits of its

range here are unclear, but it is known from Cane Creek, McDowell

County, and Green River, Henderson County), C. sp. A (Hobbs and

Peters 1977:52), C. asperimanus (in southeastern McDowell and

northeastern Cleveland counties), and C. sp. C (which is probably

118 J. E. Cooper and A. L. Braswell

throughout the system). The ubiquitous P. a. acutus may turn up in

the eastern reaches of the basin, and it is not inconceivable that

exhaustive effort could reveal C. carolinus along the border with

South Carolina. The latter species occurs in the upper Tyger River

watershed of the Broad-Congaree basin, in Greenville and Pickens

counties, South Carolina (Hobbs and Bouchard 1973:60-61). A juvenile

male crayfish (NCSM C-2229) that strongly resembles Cambarus

(Puncticambarus) spicatus Hobbs was collected in the North Pacolet

River of the lower Broad basin in Polk County, but it would be

premature to add this species to the North Carolina fauna on the basis

of this specimen. The species is known with certainty only from a

tributary of the Broad River in Fairfield and Richland counties,

South Carolina (Hobbs 1989:27).

Catawba River — This heavily impounded system rises in North

Carolina, with its headwaters on the Blue Ridge escarpment in south-

eastern Avery, Caldwell, McDowell, and Burke counties. The streams

flow into and through the Piedmont Plateau. Tributaries that head in

southern Mecklenburg and western Union counties flow independently

south-southwest into South Carolina, where they join the Catawba

River in York County. The Catawba flows south into South Carolina,

where it enters the Wateree basin of the Santee drainage.

Cambarus b. bartonii and C. asperimanus appear to be limited

to the foothills and upper Piedmont sections of the Catawba River

basin. The former may occur as far south and east as Catawba County

at the Alexander County line (Hobbs and Peters 1977:45-46), unless

the population there is revealed to be C. sp. A. Cambarus asperimanus

has been found in streams in Burke, Catawba, and McDowell counties.

Cambarus dubius has been reported from Avery County in the

upper Catawba basin (Hobbs and Peters 1977:24). Cambarus sp.

A and C. sp. C may occur throughout the system, and C. sp. B

currently is known only from Catawba County. Cambarus reduncus is

known from Gaston, Mecklenburg, and Union counties. Procambarus

a. acutus has been collected in Lincoln and Gaston counties. Hobbs

and Peters (1977:45) recorded Cambarus (Hiaticambarus) longulus

from a locality in the Catawba basin in northern Caldwell County,

but there is evidence that the specimens from that site either came

from a tributary of the nearby Yadkin-Pee Dee River, or belong to

what appears to be an undescribed species under investigation by

JEC. James (1966:13) did not show C. longulus in the Catawba basin,

and Hobbs later (1989:20) wrote that it ranges "south to the Yadkin

basin in North Carolina . . . ."

North Carolina Crayfishes 119

Yadkin-Pee Dee River — The headwaters of the Yadkin-Pee

Dee basin rise in the eastern Blue Ridge of Virginia, in western

Surry, Wilkes, and southeastern Watauga counties, North Carolina,

and in the western Piedmont in southwestern Stokes and western

Forsyth counties, North Carolina. The main trunk of the Yadkin River

begins in Watauga County east of Blowing Rock, flows southeast to

northcentral Caldwell County, then makes an abrupt turn and flows

northeast to the juncture of Surry, Stokes, Yadkin, and Forsyth

counties. There it turns south and, at the Stanly-Montgomery county

line southeast of Badin Lake, joins the Uwharrie River, a major eastern

tributary that rises in northwestern Randolph County. This merger

forms the Pee Dee River. A major western tributary, the Rocky River,

heads in southern Iredell and northern Mecklenburg counties, flows

east, and joins the Pee Dee at the Anson County line below Lake

Tillery. Several creeks that drain southcentral Union County flow

south directly into the Lynches River, South Carolina, a major

western trunk of the Pee Dee River. Southeastern tributaries of the

Pee Dee drain southern Anson and Richmond counties, flowing

directly into South Carolina to join the Pee Dee River in Chesterfield

County. The Pee Dee flows across northern South Carolina and

merges with the Little Pee Dee River at the tip of Britton Neck. The

Pee Dee then continues into Winyah Bay.

In the uppermost sections of the Yadkin-Pee Dee basin occur

C. asperimanus, C. longulus, and C. dubius. The southern limits of

their distributions are unknown, but all three appear to be restricted

to higher elevations within this basin. Specimens tentatively

identified as C. b. bartonii have also been collected in the upper

reaches of this basin. Cambarus reduncus occurs in the Piedmont,

and C. sp. C apparently occurs throughout. Hobbs and Peters (1977:

18) reported C. sp. A from Mongtomery County, collected with C. sp.

C and P. a. acutus, inadvertently placing the locality ("creek, 4.6

mi . . . W Mt Gilead on SR 73") in the Catawba River basin. Holt

and Weigl (1979:24) described the ectocommensal branchiobdellid,

Xironodrilus bashaviae, from several creeks in Forsyth County, includ-

ing "Bashavia" (Barshavia = Bechewa) Creek, with C. b. bartonii its

host. This crayfish could turn out to be C. sp. A. Procambarus a.

acutus has been collected in Anson, Cabarrus, Davidson, Montgomery,

and Union counties. Fallicambarus fodiens and C. diogenes may occur

in that part of the basin draining the southeastern Piedmont and

Sandhills.

Lumber River — To the west this Coastal Plain system heads in

the Sandhills of southern Moore, eastern Montgomery, and Richmond,

120 J. E. Cooper and A. L. Braswell

Scotland, and Hoke counties. Some of the streams drain directly into

the Little Pee Dee River just across the South Carolina line. Eastern

tributaries rise in western Bladen and Columbus counties. The Lumber

River flows primarily east and south into South Carolina, where it

joins the Little Pee Dee River on the Marion-Horry county line south

of Nichols.

The crayfish fauna of the Lumber basin consists of C. diogenes,

C. sp. C, F. fodiens, P. a. acutus, Procambarus (Ortmannicus)

ancylus Hobbs, Procambarus {Ortmannicus) blandingii (Harlan), and

P. pearsei. A locality for the latter species, given as "CAPE FEAR

BASIN. Bladen County: (2) 3.7 mi (6 km) S Dublin on St Rte 41"

(Hobbs and Peters 1977:52), actually is in the Lumber River basin.

An unidentified Procambarus, currently represented in collections

by only juveniles and one female, has been found in the Lumber

River basin in Robeson and Scotland counties. When form I

males become available, this species may prove to be Procambarus

(Ortmannicus) lepidodactylus Hobbs, which is known from the system

in South Carolina, or an undescribed species of the Pictus Group

currently known only from the Waccamaw River basin (see below).

Waccamaw River — The northernmost headwaters of the

Waccamaw River drain southcentral Bladen and northern Columbus

counties, and its eastern tributaries drain western and northern

Brunswick County. Some of the northern tributaries enter Lake

Waccamaw, the largest of the natural Carolina Bay lakes, and the

Waccamaw River flows from the south shore of the lake. East of the

lake there is an extensive system of man-made drainage canals that

"connects Honey Island Swamp (Juniper Creek tributary) with Dans

Creek of the Cape Fear drainage and Big Creek which drains into

Lake Waccamaw, . . . ." (Shute et al. 1981:21). South of the lake,

the Waccamaw River forms part of the line between western

Brunswick and southeastern Columbus counties, then flows

southwest into South Carolina and enters Winyah Bay. The Shallotte,

Lockwood Folly, and Calabash rivers, which drain most of southern

Brunswick County, are here included in this system. The Calabash

River flows west into Little River in Horry County, South Carolina,

and is connected to the Waccamaw River through the Intracoastal

Waterway.

The crayfish fauna of the Waccamaw basin consists of C.

diogenes, C. latimanus (known from two localities in Brunswick

County), F. fodiens, P. a. acutus, P. ancylus, P. blandingii, and P.

pearsei. Specimens from this basin previously assigned to P.

lepidodactylus actually represent a new species being described by

North Carolina Crayfishes 121

JEC. Current evidence indicates that C. sp. C, which occurs in the

adjacent Lumber River basin, is absent from the Waccamaw basin.

Cape Fear River — The Cape Fear River, which begins in the

Piedmont Plateau and drains the largest watershed in the State, is

confined to North Carolina. Its major northern tributary, the

Haw River, heads in northwestern Guilford, southern Rockingham,

Orange, and Durham counties. A large southern tributary, the Deep

River, begins in southwestern Guilford and eastern Randolph counties,

flows southeast to northern Moore County, then turns east and north,

forming the northwestern boundary of Lee County. The Haw and

Deep rivers then merge in eastern Chatham County at the Lee County

line, forming the Cape Fear River. A central tributary, the Rocky

River, heads in northeastern Randolph County, flows southeast

through Chatham County, and joins the Deep River at the Lee County

line. A major central tributary in the Coastal Plain, the Black River,

drains Sampson County and the western part of Duplin County.

Another Coastal Plain tributary, the South River, divides Cumberland

and Bladen counties to the west from Sampson and Pender counties to

the east. The Black River merges with the South River in southern

Sampson County at the Bladen County line, and the South enters the

Cape Fear at the line between Pender and Bladen counties. The Cape

Fear then empties into the estuary at Wilmington, New Hanover

County.

Until recently, the endemic North Carolina crayfish, C. catagius,

was known only from its type locality, burrows in a lawn at East

Whittington Street in the southeastern section of Greensboro,

Guilford County (Hobbs and Perkins 1967:145), which is in the Haw

River subdrainage. Other colonies of this primary burrower have now

been discovered, and efforts are being made to more accurately

determine its range (Davis 1992:29). Cambarus reduncus is restricted

to the Piedmont and Fall Line zone, while C. latimanus, C. diogenes,

and C. sp. C are found in both the Piedmont and Coastal Plain. Procambarus

a. acutus appears to be common throughout the system, even into its

headwaters. Faxon's (1914:367) report of "Cambarus blandingii" from

Reedy Creek, near Greensboro, must have been based on P. a. acutus,

which D. G. Cooper collected from Lake Reidsville and a small lake

south of Reidsville, Rockingham County, in the Haw River subdrainage.

Fallicambarus fodiens apparently has made inroads into the Piedmont

of the Cape Fear basin; Hobbs and Peters (1977:46) recorded it from

the Rocky River in Chatham County. Procambarus ancylus is known

from White and Singletary lakes, Bladen County (Hobbs 1958:167),

and has been collected from a borrow pit near Colly Creek (NCSM

122 J. E. Cooper and A. L. Braswell

C-1949), which has a tributary stream entering it from Singletary

Lake. Procambarus pearsei is known from as far north in the system

as northern Sampson County, and might occur in extreme southwestern

Johnston County. Its type locality, a pond and ditch on NC 22 south

of Fayetteville, Cumberland County (Creaser 1934:4), is in this river

basin. Procambarus blandingii, which occurs in the Lumber and Waccamaw

basins, appears to be absent from the lower Cape Fear, but this needs

investigation.

The northern tributaries of the Northeast Cape Fear River originate

in the Coastal Plain in southern Wayne and northeastern Sampson

counties. The system drains almost all of Duplin, most of Pender,

western Onslow, and northern New Hanover counties. The Northeast

Cape Fear generally is considered an eastern trunk of the Cape Fear

River, but it flows into saline waters at Wilmington, independent of

the Cape Fear. As alluded to in the previous discussion of the Northeast

Cape Fear as a hydrologically and faunistically autonomous unit, there

are differences in the crayfish faunas of the two systems. Since the

Northeast Cape Fear is confined to the Coastal Plain, it is not surprising

that it does not harbor C. catagius or C. reduncus, both Piedmont

species. It does have P. plumimanus, however, which is absent from

the Cape Fear but present in the New (White Oak), and lacks P.

ancylus and P. pearsei, both of which occur in the Cape Fear, Lumber,

and Waccamaw. The rest of the crayfish fauna of the Northeast Cape

Fear is that of the coastal Cape Fear, i.e., C. latimanus, C. diogenes,

C. sp. C, F. fodiens, and P. a. acutus.

New {White Oak) River — The major hydrologic units of the New

(White Oak) drainage basin are the autonomous New, White Oak,

Newport, and North rivers and their tributaries. All are small and all

empty into saline coastal waters behind barrier islands in Onslow

Bay, and Bogue and Back sounds. The New River drains the center of

Onslow County, and extreme eastern Pender and New Hanover

counties. The White Oak drains eastern Onslow County and forms

part of the boundary between southeastern Jones and eastern Onslow

counties. Included in the White Oak unit are Catfish and Great lakes

of Croatan National Forest, southwestern Craven County. The Newport

and New rivers both drain southern Carteret County, and both, like

the White Oak, are artificially connected with the lower Neuse River

basin to the north (see the P. plumimanus account).

The crayfish fauna of the New (White Oak) basin consists of

C. diogenes, C. latimanus, F. fodiens, P. a. acutus, P. plumimanus,

and probably C. sp. C.

North Carolina Crayfishes 123

Neuse River — The Neuse is a second major drainage system

that heads and debouches in North Carolina (for a brief description

see Cooper and Ashton 1985:6-7). It includes Long, Little, and Ellis

lakes of Croatan National Forest, Craven County. Ellis Lake has a

direct connection with Southwest Prong Slocum Creek.

Cambarus reduncus is confined to the Piedmont Plateau and

Fall Line zone of this basin, where the southeasternmost known

localities are in central Wake County. Cambarus latimanus, C. diogenes,

C. sp. C, and P. a. acutus occur throughout the system. Procambarus

medialis, F. fodiens, and O. sp. A are essentially Coastal Plain species,

with F. fodiens and O. sp. A both edging into the Fall Line zone at

least as far west as southern Wake County. Two localities for P.

plumimanus, including the type locality, are known in the lower Neuse

basin. The listing by Harris (1903:56) of C. b. bartonii in the Neuse

was in error, and was probably based on Faxon's (1885:61) erroneous

report of the species from Kinston, Lenoir County, which also was

repeated by Ortmann (1931:131).

Tar-Pamlico River — Like the Cape Fear and Neuse rivers, the

Tar-Pamlico is confined to North Carolina (for a brief description see

Cooper and Ashton 1985:6-9). Lake Mattamuskeet, part of this system

in Hyde County, connects with the Alligator River of the Pasquotank

basin through drainage canals and the Intracoastal Waterway, which

also forms a direct connection between the Alligator and Pungo rivers.

Drainage canals also connect Pungo Lake of the Tar-Pamlico basin,

Washington and Hyde counties, with canals from New (Alligator)

Lake in the Pasquotank drainage of Hyde County.

The crayfish fauna of the Tar-Pamlico is the same as that of

the Neuse basin, except for the apparent expansion into the latter of

P. plumimanus. During the Pleistocene, these drainages "would have

been conjoined to form the Greater Pamlico River" (Lachner and

Jenkins 1971:62). In the Tar-Pamlico, however, P. medialis appears

to be limited to the immediate area of the type locality, a pool in

a roadside ditch, 0.6 mi (1 km) south of Scotland Neck, on US 258, in

southeastern Halifax County (Hobbs 1975:13). In the Neuse basin its

known range is more extensive, occupying an area from Johnston

County downriver to Lenoir and Pitt counties. In addition, the Tar-

Pamlico range of Orconectes sp. A extends from headwaters in

Granville County to eastern Pitt County, whereas in the Neuse, despite

intensive sampling, this species has not been found any farther

upriver than southern Wake County. The report by Faxon (1914:367)

of "Cambarus blandingii" from Lake Mattamuskeet was undoubtedly

based upon P. a. acutus.

124 J. E. Cooper and A. L. Braswell

Roanoke River — The Roanoke is another heavily impounded

river. Its major western tributary, the Dan River, begins in the Blue

Ridge in Virginia, and in northeastern Surry, Stokes, and Forsyth

counties, North Carolina. The Dan flows essentially east and north,

moving in and out of North Carolina in northern Rockingham and

Caswell counties, and acquiring tributaries in both. It joins the

Roanoke at Buggs Island Reservoir in Virginia. The Roanoke then

flows southeast into North Carolina, adding tributaries in northern

Granville and Vance counties and feeding into John H. Kerr Reservoir

and Lake Gaston. The river flows southeast from Lake Gaston, drains

southwestern Northampton, northern Halifax, and most of Bertie and

Martin counties, then empties into the western arm of Albemarle

Sound at Batchelor Bay.

Cambarus longulus and C. b. bartonii occur in the upper Dan

River watershed, where C. longulus is known from no farther east

than Rockingham County, and the eastern limits of the range of C. b.

bartonii are unknown. Cambarus sp. C apparently occurs throughout

the system. Cambarus diogenes and F. fodiens occur in the Coastal

Plain, but encroach on the Piedmont Plateau, where the western limits

of their ranges are uncertain. Procambarus a. acutus is found in all of

the Coastal Plain, and as far west in the Piedmont as eastern Caswell

County. Orconectes virginiensis is known from Bertie, Halifax, and

Martin counties.

Chowan River — The Chowan River basin is confined to the

northeastern Coastal Plain, rising at the confluence of the Nottoway

and Blackwater rivers just south of the Virginia state line. The southeast-

flowing Meherrin River heads in Virginia, enters North Carolina to

form part of the northern boundary between Northampton and Hertford

counties, then turns east across Hertford County to join the Chowan

at the Gates County line. The system drains northern and eastern

Northampton, Hertford, and most of southern and western Gates,

western Chowan, and northeastern Bertie counties, then empties

into the western arm of Albemarle Sound.

The crayfishes of the Chowan basin are C. diogenes, F. fodiens,

P. a. acutus, O. virginiensis, and possibly C. sp. C. The listing by

Harris (1903:53) of C. b. bartonii in the Chowan basin (in Virginia)

was in error, and was probably based on Faxon's (1885:61) erroneous

report of the species from Southampton and Lunenburg counties,

which also was repeated by Ortmann (1931:131)

Pasquotank River — North of Albemarle Sound, the streams

that form the northern part of this Coastal Plain system rise in the

southeastern tip of Virginia and in eastern Gates County, North

Carolina. This part of the basin includes the Pasquotank River that

North Carolina Crayfishes 125

drains northeastern Gates, northern Pasquotank, and the northern

half of Camden counties; the Perquimans River of Perquimans County;

the Little River that rises in the Dismal Swamp in central Pasquotank

County and forms part of the lower Pasquotank-Perquimans county

line; the North River that rises in the Dismal Swamp in Currituck and

Camden counties; the Northwest River, which begins in Norfolk

County, Virginia, and flows southeast into Tull Bay, Currituck County;

and the North Landing River, which begins in Princess Anne County,

Virginia, flows south into Currituck Sound, and connects with the

lower James River of Virginia via the Intracoastal Waterway. All

these rivers except the Northwest and North Landing flow into

Albemarle Sound.

South of Albemarle Sound the Pasquotank system includes the

north-flowing Alligator River, which drains parts of northern Hyde,

much of Tyrrell, and most of Dare counties; the Scuppernong River

that drains eastern Washington and northwestern Tyrrell counties;

and Phelps and New (Alligator) lakes. Both an immense system of

man-made drainage canals in the Dismal Swamp west of Phelps and

Pungo lakes, and the Intracoastal Waterway, link parts of the Pasquotank

basin with the Pungo River, Lake Mattamuskeet, and other parts of

the Tar-Pamlico basin in Beaufort and Hyde counties.

The crayfish fauna of the Pasquotank basin consists of C.

diogenes, F. fodiens, and P. a. acutus. Fallicambarus fodiens has

been found across Croatan Sound, near Manteo on Roanoke Island,

Dare County.

CRAYFISH DISTRIBUTIONS BY PHYSIOGRAPHIC PROVINCES

Cambarus is the dominant crayfish genus in North Carolina,

with 18 described species (not including C. " acuminatus") and at

least three known but undescribed species. Nine of these species are

limited to the Blue Ridge, three are limited to the Piedmont

Plateau, and two overlap the Piedmont and Coastal Plain (one of

these two also may be found in all three provinces). Distributions of

the taxa now subsumed under Cambarus sp. C, which includes

C. "acuminatus" will not be clarified until this complex has been

diagnosed. Suffice it to say that they occur in all three physiographic

provinces. The sole North Carolina species of Fallicambarus is

primarily Coastal Plain in distribution, but also has invaded the eastern

Piedmont Plateau. One of the two described Orconectes in North

Carolina occurs only in the Blue Ridge, as does an apparently

undescribed member of the genus. The other described Orconectes is

limited to the northeastern Coastal Plain, and a second undescribed

126

J. E. Cooper and A. L. Braswell

Table 2. Summary of crayfish distributions in major physiographic provinces

of North Carolina. X = present.

1 Known from east of the Blue Ridge in Avery, Surry, and Wilkes counties, but

eastern limits of range unknown.

2 Includes C. (P.) acuminatus (s. I.).

North Carolina Crayfishes 127

species of the genus is essentially Coastal Plain but occurs also at

the eastern edge of the Piedmont Plateau. Genus Procambarus, with

six, and probably seven, described native species, dominates the

Coastal Plain. Six of these species are limited to this province, but

the seventh also occurs deeply into the Piedmont. Table 2 is a summary

of species distributions by physiographic provinces.

ACKNOWLEDGMENTS— -We express our sincerest thanks to all

those collectors mentioned in this paper, without whose efforts we

would know considerably less about North Carolina crayfishes than

we do. Gary W. Woodyard provided the blue C. latimanus and helped

in other ways. Joseph F. Fitzpatrick, Jr. and Martha R. Cooper shared,

as usual, their knowledge and libraries. Dr. Fitzpatrick and an anonymous

reviewer provided invaluable comments on the manuscript. JEC is

also indebted to John E. Cooper, Jr., Martha R. Cooper, Gladys Leake,

Jesse Perry, Lynn Ferguson, John R. Holsinger, William J. Rishel,

Patricia S. and Ray E. Ashton, Jr., his co-author, and especially Don

Howard, without whose assistance this paper could never have been

completed.

Finally, we express our deepest gratitude to and for the late

Horton H. Hobbs, Jr., whose death was a great personal tragedy and

an incalculable loss to decapod research. His kindness and generosity,

his knowledge and insights, and his total integrity, shall never be

forgotten.

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132 J. E. Cooper and A. L. Braswell

ADDENDUM

After this paper went to press, V. Schneider and D. R. Lenat

collected the first Cambarus spicatus adults from North Carolina:

Cleveland Co., First Broad R at SR 1530, 6.1 air mi (9.8 air km)

WSW cntr Casar; 2 S II, 3 9 (NCSM C-2487), 20 Jun 1995.

Received 29 September 1994

Accepted 12 January 1995

133

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21:69-72) have requested that the following corrections be published:

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134

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BRIMLEYANA NO. 22, JUNE 1995

CONTENTS

Aspects of the Feeding Ecology of the Little Grass Frog, Pseudacris ocularis

(Anura: Hylidae). Jeremy L. Marshall and Carlos D. Camp

Corrections of Records of Occurrence of Peromyscus polionotus (Wagner) and

P. gossypinus (LeConte) (Rodentia: Muridae) in the Blue Ridge Province

of Georgia. Joshua Laerm and James L. Boone

The Masked Shrew, Sorex cinereus (Insectivora: Soricidae), and the Red-backed Vole,

Clethrionomys gapperi (Rodentia: Muridae), in the Blue Ridge Province of

South Carolina. Joshua Laerm, Eric Brown, Michael A. Menzel, Amanda

Wotjalik, William Mark Ford, and Mary Strayer 15

Rediscovery of the Aquatic Gastropod Helisoma eucosmium (Bartsch, 1908),

(Basommatophora: Planorbidae). William F. Adams and Susan G. Brady 23

Effects of a Clearcut on the Herpetofauna of a South Carolina Bottomland Swamp.

Jospeh P. Phelps and Richard A. Lancia 31

First Record of the Water Shrew, Sorex palustris Richardson (Insectivora: Soricidae),

in Georgia with Comments on its Distribution and Status in the Southern

Appalachians. Joshua Laerm, Charles H. Wharton, and William Mark Ford 47

Florida Manatees, Trichechus manatus (Sirenia: Trichechidae), in North Carolina

1919-1994. Frank J. Schwartz 53

Recovery of the Cave Crayfish (Decapoda: Cambridae) Population in Peacock

Springs, Florida? W. J. Streever 61

Premolar Cementum and Noncementum Lengths As Potential Indicators of Age

for Beavers, Castor canadensis (Rodentia: Castoridae). Allan E. Houston

and Michael R. Pelton 67

Record Clutch Size for Chelydra serpentina (Testudines: Chelydridae) in Virginia.

Joseph C. Mitchell and Michael C. Odom 73

New Distributional Records for the Star-nosed Mole, Condylura cristata

(Insectivora: Talpidae), in North Carolina, with Comments on its

Occurrence in the Piedmont Region. Jeffrey C. Beane 77

Observations on North Carolina Crayfishes (Decapoda: Cambaridae).

John E. Cooper and Alvin L. Braswell 87

Miscellany 133